ID: 

0 


lilMiiii: 


i 


ill 


'■ill 


'il! 


iillili^ 


illiiiliiil 


^!ili^     ^ilii 


(PKINS  HOSPITAL  REPOR 


THE  JOHNS  HOPKINS  HOSPITAL  REPORTS 
MONOGRAPHS.     NEW  SERIES  No.  VI 


THE  NUCLEI  TUBERIS  LATERALES 

AND  THE  SO-CALLED  GANGLION 

OPTICUM  BASALE 

BY 
EDWARD  F.ImALONE 

(From  the  Anatomical  Laboratory  of  the  University  of  Cincinnati) 
Illustrated— 15  Plates 


BALTIMORE 

THE  JOHNS  HOPKINS  PRESS 

1914 

fCopyright,  1914.  by  The  Johns  Hopkiiu  Press] 


«!• 


CONTENTS. 

PAGE 

I.  Introduction 1 

II.  Material  3 

III.  Pars  Mammillaris  Hypothalami 4 

IV.  Ganglion  Opticum  Basale 7 

A.  Location  and  extent  of  tlie  ganglion  opticum  basale 

1.  Man 8 

2.  Macacus  rhesus  11 

3.  Lemur  rufus  11 

4.  Cat  12 

B.  Cell  type  of  the  basal  optic  ganglion 13 

C.  Separation  of  the  basal  optic  ganglion  from  surround- 

ing cell  groups  through  differences  in  cell  char- 
acter    14 

V.  Nuclei  Tuberis  Laterales 18 

A.  Location  and  extent  of  the  nuclei  tuberis  laterales 

1.  Man 19 

2.  Macacus  rhesus   21 

B.  Separation  of  the  nuclei  tuberis  laterales  from  sur- 

rounding cell  groups  through  diiterences  in  cell 

character 21 

VI.  Relationship  of  the  Various  Cell  Groups  to  One  Another 23 

VII.  Conclusion    27 

VIII.  Literature  31 

IX.  Bibliography    38 

X.  Explanation  of  Illustrations  39 


THE  NUCLEI  TUBEEIS  LATERALES  AND  THE  SO-CALLED 
GANGLION  OPTICUM  BASALE. 

By  EDWARD  F.  MALONE, 

(From  the  Anatomical  Laboratory  of  the  UniverHty  of  Cincinnati.) 

There  is  perhaps  no  region  of  the  mammalian  brain  concerning 
whose  cell  groups  less  is  known  than  that  portion  of  the  telencephalon 
known  as  the  pars  optica  hypothalami.  The  cell  groups  which  consti- 
tute the  subject  of  this  article  are  so  vague  and  so  hopelessly  confused 
in  the  literature  that  it  is  not  even  possible  to  determine  whether  the 
names  basal  optic  ganglion  and  nuclei  tuberis  should  be  applied  to 
different  cell  groups  or  to  different  portions  of  the  same  cell  group. 
In  this  article  it  will  be  shown  that  these  names  should  be  applied  to 
entirely  different  cell  groups ;  moreover  the  location  and  extent  of  the 
two  cell  groups  will  be  described  in  different  mammals,  together  with  a 
consideration  of  their  relation  to  surrounding  groups  of  cells,  and  it 
will  be  shown  that  these  two  cell  groups  under  consideration  are  com- 
posed of  cells  of  radically  different  character  through  which  each  group 
may  be  readily  distinguished  from  the  other  and  from  the  surrounding 
cell  groups.  In  addition  the  cell  groups  of  the  pars  mammillaris  of  the 
hypothalamus  will  be  described. 

When  we  consider  the  methods  employed  by  most  of  those  who  study 
the  anatomy  of  the  mammalian  brain  perhaps  it  will  not  be  surprising 
to  realize  that  the  best  description  of  the  basal  optic  ganglion  and  the 
nuclei  tuberis  is  that  of  Kolliker  published  in  1896,  however  faulty  and 
confusing  this  description  may  be.  The  reason  for  this  lack -of  infor- 
mation concerning  these  cell  groups  is  that  most  workers  are  interested 
in  the  course  of  fiber  tracts,  and  their  interest  in  the  structure  of  the 
nervous  system  is  for  the  most  part  limited  to  the  largely  mechanical 
subdivisions  revealed  by  fiber  stains ;  in  such  preparations  the  course  of 
fiber  systems  may  often  be  followed,  or  at  least  they  reveal  certain 
architectural  differences  of  various  regions  which  serve  the  purpose  of 
orientation.  When  preparations  in  which  the  cells  are  stained  are 
studied  these  preparations  are  for  the  most  part  regarded  as  showing 
merely  the  negative  picture  of  the  fiber  preparations,  and  the  same 


2  Edward  F.  M alone. 

topograpliical  regions  are  observed,  the  cells  being  stained  instead  of 
the  fiber;;.  When  the  cell  character  is  noted  it  is  done  usually  in  a 
superficial  manner  and  principally  for  the  purpose  of  orientation,  and 
with  no  purpose  of  bringing  the  cell  character  into  relation  with  a 
definite  function. 

The  same  lack  of  attention  to  the  cell  tyjie  is  apparent  even  in  the 
work  of  the  relatively  few  investigators  who  employ  the  experimental 
methods  of  Xissl  or  von  Gudden.  The  following  instance  is  a  good 
illustration  of  the  value  of  carefully  noting  the  cell  character  of  various 
cell  groups.  In  his  excellent  study  of  the  dorsal  (sympathetic)  nucleus 
of  the  vagus  nerve  Molhant  had  shown  that  all  the  cells  of  this  cell 
column  give  origin  to  all  of  the  vagus  fibers  which  supply  smooth 
muscle  and  heart  muscle,  and  that  certain  portions  of  this  cell  column 
supply  smooth  muscle  while  a  definite  portion  supplies  heart  muscle, 
but  he  did  not  attempt  to  show  that  a  definite  type  of  cell  was  involved 
in  the  innervation  of  each  of  these  two  types  of  muscle ;  after  studying 
the  vagus  sympathetic  nucleus  I  was  able  to  show  that  the  portions  of 
the  nucleus  which  supply  smooth  muscle  and  heart  muscle  may  be 
readily  distinguished  by  the  fact  that  they  are  composed  of  cells  of 
difi'erent  types,  and  that  just  as  heart  muscle  is  histologically  inter- 
mediate between  smooth  muscle  and  striated  muscle,  just  so  the  cells 
of  the  vagiis  sympathetic  nucleus  which  supply  heart  muscle  are  of  a 
histological  character  intermediate  between  that  of  the  cells  which 
supply  smooth  muscle  and  striated  muscle.  Such  an  observation  of  the 
relation  of  cell  type  to  cell  function  enables  us  to  locate  accurately  a 
functional  center  even  though  its  cells  be  mixed  with  those  having  a 
different  function,  and  homologous  centers  may  be  recognized  in 
different  animal  fonns;  at  the  same  time  it  emphasizes  the  necessity 
of  carefully  noting  the  cell  character  of  each  cell  group,  so  that  even  if 
the  function  is  imknown  the  presence  of  a  definite  cell  type  will  offer  a 
problem  for  future  experimental  work,  and  in  the  meanwhile  preclude 
the  possibility  of  confusing  this  cell  group  with  surrounding  cells. 
These  differences  in  cell  type  are  of  course  most  definite  in  the  higher 
mammals  and  especially  in  man,  where  the  various  cell  groups  are 
highly  specialized. 

The  foregoing  consideration  of  the  failure  of  workers  on  the  anatomy 
of  the  mammalian  brain  to  attach  sufficient  importance  to  the  different 
types  of  cells  explains  why  such  characteristic  cell  groups  as  the  basal 
optic  ganglion  and  the  nuclei  tuberis  have  not  been  clearly  separated 


Nuclei  Tuberis  Laterales  and  the  Ganglion  Opticum  Basale.      3 

from  one  another  and  from  the  surrounding  cell  groups.  The  unfor- 
tunate condition  which  here  exists  is  present  also  in  many  other  regions 
of  the  nervous  system,  and  I  have  considered  the  tendency  of  the  anatom- 
ical work  on  the  mammalian  brain  somewhat  at  length  not  merely  as 
an  interesting  explanation  for  our  present  want  of  definite  information, 
but  because  this  tendency  contains  a  serious  defect  whose  results  are 
most  evident  and  far  reaching,  a  defect  which  should  be  corrected. 

The  tendency  of  neurological  workers  to  attach  little  importance  to 
differences  in  cell  character  I  have  criticised,  not  only  at  this  point  but 
also  later  in  this  article.  On  the  other  hand  I  have  no  intention  what- 
ever of  disparaging  the  ability  of  other  authors,  nor  am  I  unapprecia- 
tive  of  the  excellence,  in  many  cases,  of  their  work  and  of  the  valuable 
results  which  they  have  contributed.  My  criticism  is  aimed  exclusively 
at  the  almost  universal  belief  that  the  various  differences  in  cell  type 
in  the  nervous  system  are,  after  all,  of  no  great  importance,  and  that 
We  should  reserve  our  efforts  to  discover  the  connections  of  various 
portions  of  the  nervous  system.  It  would  be  unfair  to  blame  authors  for 
being  influenced  by  a  belief  so  general;  moreover  it  is  only  compara- 
tively recently  that  we  have  begun  to  employ  methods  capable  of  reveal- 
ing the  striking  differences  in  cell  character,  and  only  in  recent  years 
has  the  development  of  cytology  given  an  added  stimulus  to  the  careful 
study  of  the  nerve  cell.  Above  all  I  wish  to  disclaim  any  belief  that  the 
method  herein  employed — that  of  setting  aside  groups  of  cells  of  an 
identical  type  and  of  comparing  such  cell  groups  and  of  attempting  to 
discover  the  correlation  between  cell  tj'pe  and  cell  function — is  of  any 
more  value  than  many  other  methods  of  investigation.  I  advocate  it  not 
to  displace  other  methods,  but  to  complement  them  and  add  to  their 
usefulness. 

Material. 

The  material  studied  consists  of  the  following  complete  series :  four  of 
man,  one  of  macacus  rhesus,  two  of  lemur  rufus,  and  three  of  the  cat. 
The  tissue  was  in  each  case  fixed  in  95  per  cent  alcohol,  and  after  dehy- 
dration and  treatment  with  chloroform  embedded  in  paraffin.  The 
sections  were  stained  in  a  one  per  cent  aqueous  solution  of  toluidin  blue 
(Griibler),  differentiated  in  95  per  cent  alcohol,  dehydrated  in  absolute 
alcohol,  cleared  in  xylol,  and  mounted  in  Canada  balsam. 


4  Edward  F.  Malone. 

Pars  Mamsiillaris  Hypothal.\mi. 

Before  proceeding  to  describe  the  cell  groups  of  the  pars  optica  hypo- 
thalami (telencephalon)  it  will  be  necessary  to  consider  those  of  the 
pars  mammillaris  hypothalami  (diencephalon),  since  the  cell  groups 
of  these  two  subdivisions  of  the  hypothalamus  are  most  intimately 
related  and  cannot  be  intelligently  studied  separately.  In  1910 
appeared  my  monograph  "  Uber  die  Kerne  des  mensehlichen  Dienceph- 
alon "  in  which  the  cell  groups  of  the  pars  mammillaris  hypothalami 
of  man  were  described,  together  with  a  brief  consideration  of  certain 
closely  related  cell  groups  of  the  pars  optica.  Since  the  publication  of 
this  monograph  I  have  studied  both  portions  of  the  hypothalamus  in 
the  monkey,  lemur  and  cat,  and  have  compared  the  cell  groups  observed 
in  these  animals  with  homologous  groups  found  in  man.  The  following 
description  of  the  cell  groups  of  the  pars  mammillaris  of  the  monkey, 
lemur  and  cat  is  accordingly  original  and  I  should  like  to  call  especial 
attention  to  the  illustrations  which  clearly  show  for  the  first  time  the 
different  cell  groups  in  macacus  rhesus  (Figs.  19  to  25).  In  Series 
D  of  man  these  cell  groups  are  also  well  sIiowti  (Figs.  11  to  18).  The 
other  series  do  not  show  much  of  the  pars  mammillaris,  since  it  did  not 
appear  advisable  to  add  many  illustrations  to  show  exclusively  portions 
of  the  pars  mammillaris  which  are  not  intimately  related  to  the  pars 
optica.  In  the  series  of  macacus  and  the  Series  D  of  man  the  plane  of 
section  is  such  as  to  include  this  caudal  portion  of  the  pars  mammil- 
laris. Moreover  the  dorsal  portion  of  the  pars  mammillaris  has  not 
been  illustrated,  not  only  because  it  has  no  close  relation  to  the  pars 
optica  but  also  because  the  thalaanus  of  the  three  lower  forms  must 
first  be  studied.  Certain  types  of  cells  also  which  on  account  of  their 
location  could  not  be  confused  with  the  cells  of  the  pars  optica  have  not 
been  illustrated.  Accordingly  it  appears  best  to  defer  such  numerous 
and  ekborate  illustrations  until  the  whole  diencephalon  can  be  included, 
and  to  limit  the  illustrations  of  the  present  article  to  those  portions  of 
the  pars  mammillaris  which  are  closely  related  to  the  pars  optica. 

In  studying  the  anatomy  of  the  cell  groups  of  the  diencephalon 
nowhere  are  more  satisfactory  results  obtained  than  in  the  hypothal- 
aniu.s  (pars  mammillaris).  The  great  difficulties  encountered  in 
attempting  to  homologize  the  cell  groups  of  the  thalamus  of  different 
mammals  disappears  when  we  reach  the  hypothalamus.  This  is  due  to 
the  fact  that  the  thalamus  is  phylogenetically  a  much  more  recent 


Nuclei  Tuheris  Lateral es  and  the  Ganglion  Opticum  Basale.       5 

portion  of  the  brain  and  its  development  is  largely  dependent  upon  that 
of  the  rapidly  developing  pallium.  So  small  is  the  difference  in  the  cell 
groups  of  the  hypothalamus  of  the  various  mammals  studied  that  for 
the  purposes  of  this  article  one  description  will  suffice  for  all.  The  pars 
mammillaris  hypothalami  is  divided  into  the  following  primary  nuclei 
(groups  of  cells  having  an  identical  histological  character)  : 

1.  Ganglion  mediale  corporis  mammillaris. 

8.  Nucleus  intercalatus  corporis  mammillaris. 

3.  Nucleus  tubero-mammillaris  (nucleus  mammillo-infundibularis). 

4.  Nucleus  paraventricularis  hypothalami. 

5.  Corpus  hypothalamicum   (Luysii). 

6.  Substantia  reticularis  liypotlialami. 

7.  Substantia  grisea  ventriculi  tertii. 

At  this  point  it  is  advisable  to  read  the  explanation  of  the  various 
kinds  of  illustrations  as  given  on  pp.  39-40 ;  otherwise  the  figures,  some 
of  which  are  immediately  to  be  referred  to,  might  not  be  correctly  inter- 
preted. 

1.  Gfniglion  mediale  corporis  mammillaris. 

This  cell  group  constitutes  the  greater  portion  of  the  mammillary 
body,  and  extends  further  caudally  than  the  other  two  groups.  It  is 
shown  in  the  series  of  macacus  (Figs.  19  and  20)  and  in  the  Series  D 
of  man  (Fig.  11)  ;  no  description  of  its  relations  will  be  given.  Con- 
cerning its  cells  it  will  suffice  to  state  that  they  are  of  a  type  readily 
distinguishable  from  the  other  two  groups  of  the  corpus  mammillare. 

2.' Nucleus  intercalatus  corporis  mammillaris. 
This  is  a  small  circumscribed  cell  group,  described  by  me  in  1910, 
which  lies  between  the  medial  and  so-called  lateral  ganglion  of  the 
mammillary  body.  Its  location  may  be  seen  in  the  same  figures  that 
show  the  medial  ganglion  (see  above).  Later  Friedemann  found  this 
group  in  cercopithecus  and  adopted  the  name  here  employed.  The  cells 
of  the  nucleus  intercalatus  are  readily  distinguished  from  those  of  the 
two  other  groups  of  the  mammillar}'  body  through  their  characteristic 
type;  especially  in  man  they  have  the  relatively  large  and  discrete 
Nissl  granules  characteristic  of  efferent  nerve  cells.  This  cell  group 
should  not  be  confused  with  groups  which  are  occasionally  mechanically 
split  off  from  the  medial  ganglion  by  fiber  miasses ;  the  difference  in 
cell  type  renders  this  distinction  easy. 


6  Edward  F.  Malone. 

S.  Nucleus  tubero-mammilla/ris. 

The  cells  of  the  so-called  lateral  ganglion  of  the  corpus  mammillare 
may  be  readily  followed  orally  and  laterally  into  the  angle  between  the 
pes  pedunculi  and  the  tractus  opticus,  whereas  orally  and  dorsally  they 
may  be  followed  through  many  sections  accompanying  the  tractus 
thalamo-mammillaris  and  the  columna  fornicis.  To  this  complex  of 
cells  having  an  identical  histological  character  I  gave  (1910)  the  name 
"  nucleus  mammillo-infundibularis,"  a  name  which  Friedemann 
(1911)  has  adopted  in  his  article  on  the  diencephalon  of  cercopithecus. 
It  is  evident  that  this  name  is  unsuitable,  since  the  cell  group  is  not 
situated  in  the  infundibulum,  and  I  have  therefore  changed  it  to 
"  nucleus  tubero-mammillaris."  The  nucleus  tubero-mammillaris  is 
shown  in  all  the  series  illustrated  and  its  relation  to  the  basal  optic 
ganglion  and  the  nuclei  tuberis  may  be  readily  seen.  (Since  the  figures 
were  drawn  and  labeled  before  I  decided  to  change  the  name  of  this 
nucleus,  it  appears  on  tlie  outline  drawings  as  "•  nucleus  mammillo-in- 
fundibularis," but  wherever  space  has  permitted  the  new  name  has 
been  added  in  parentheses.  The  explanation  of  each  plate  is  so  ar- 
ranged as  to  remove  any  possibility  of  confusion.)  The  character  of  its 
cells  in  the  different  forms  is  illustrated  in  Figs.  41,  47,  52  and  57. 
The  relation  of  this  nucleus  to  the  basal  optic  ganglion  will  be  discussed 
later.  It  should  be  noted  that  a  portion  of  this  nucleus  is  situated  in 
the  pars  optica. 

Jf.  Nucleus  paraveniricularis  hypothalami. 

This  characteristic  column  of  cells  is  situated  partly  in  the  pars 
mammillaris  but  principally  in  the  pars  optica.  Its  location  is  shown 
in  all  the  series,  and  the  character  of  its  cells  in  Figs.  43,  49,  54  and 
58.  In  man  this  nucleus  was  first  described  by  me  in  1910,  and  the 
fact  that  such  a  striking  cell  group  had  not  been  previously  described 
in  man  is  probably  due  to  the  almost  exclusive  use  of  fiber  stains.  In 
1911  Friedemann  found  this  cell  group  in  cercopithecus.  It  is  certainly 
homologous  with  the  group  described  by  Cajal  in  rodents  under  the 
name  of  "  niicleo  subventricular,"'  and  ^^^th  that  described  by  Ziehen 
in  marsupials  as  "  Nucleus  subcommissuralis."  Homologous  groups 
are  also  possibly  found  in  vertebrates  even  as  low  as  the  fishes.  The 
nucleus  paraventricularis  is  intimately  related  to  the  basal  optic  gang- 
lion and  also  to  the  nucleus  tubero-mammillaris  as  tliese  three  nuclei 


Nuclei  Tuheris  Laterales  and  the  Ganglion  Opticum  Basale.       1 

appear  to  be  components  of  one  large  cell  complex,  and  while  differing 
from  one  another,  differ  much  more  from  the  surrounding  cells.  These 
relations  will  be  discussed  later  in  detail. 

5.  Corpus  hypothulamicum  (Luysii) 

This  is  such  a  compact,  isolated  cell  mass,  and  is  situated  at  such  a 
distance  from  the  basal  optic  ganglion  and  the  nuclei  tuberis  that  no 
description  is  necessary ;  it  has  been  included  in  the  illustrations,  how- 
ever, for  the  purpose  of  orientation. 

6.  Substantia  reticularis  hypothalami. 

This  group  of  cells  on  account  of  its  position  need  not  be  considered 
in  this  article.  A  description  will  be  found  in  my  monograph  on  the 
diencephalon.  Many  of  its  cells  are  of  a  motor  type  of  structure,  and 
these  together  with  the  cells  of  the  nucleus  intercalatus  of  the  corpus 
mammillare  are  the  only  cells  of  the  entire  diencephalon  which  show 
the  histological  character  peculiar  to  motor  cells. 

7.  Substantia  grisea  ventricuK  tertii. 

This  cell  mass  is  shown  in  all  series  and  the  cell  type  in  Figs.  40,  46, 
51  and  56.  The  greater  portion  lies  in  the  pars  optica,  and  it  should 
be  noted  that  dorsally  and  laterally  the  cells  are  less  densely  packed 
than  ventrally  and  medially.  The  substantia  grisea  is  closely  related 
to  the  nuclei  tuberis,  and  this  relation  will  be  fully  discussed  in  connec- 
tion with  the  description  of  the  nuclei  tuberis. 

If  we  now  observe  in  the  illustrations  the  location  and  extent  of  the 
different  cell  groups  previously  described  and  note  also  the  character  of 
their  cells  in  the  different  mammals,  we  shall  be  prepared  to  understand 
the  following  description  of  the  basal  optic  ganglion  and  the  nuclei 
tuberis  and  the  consideration  of  the  relations  of  the  latter  two  nuclei  to 
the  fonner. 

Ganglion  Opticdm  Basale. 

The  name  ganglion  opticum  basale  was  given  with  the  intention  of 
implying  a  function  which  this  cell  group  almost  certainly  does  not 
possess ;  on  the  other  hand  this  name  effectively  distinguishes  this  cell 
group  from  the  nuclei  tuberis  with  which  it  has  been  confused.  Since 
the  so-called  basal  optic  ganglion  is  such  a  characteristic  cell  group,  and 


8  Edward  F.  }[alone. 

since  the  homologous  groups  in  various  mammals  are  so  constant  both 
as  to  location,  extent  and  cell  character,  it  appears  highly  probable  that 
we  shall  not  have  to  wait  long  before  this  name  may  be  abandoned  for 
one  which  will  indicate  its  true  function,  and  until  then  it  seems  better 
not  to  introduce  a  temporary  name  implying  merely  some  morpholog- 
ical character.  In  arriving  at  the  function  of  a  cell  group  the  first 
essential  (and  probably  the  most  important  of  all)  is  to  clearly  dis- 
tinguish it  from  the  surrounding  cell  groups  and  to  study  carefully 
its  relations  to  such  cell  groups ;  in  this  article  I  shall  endeavor  by 
means  of  illustrations  and  description  to  make  the  resulting  picture 
of  the  so-called  basal  optic  ganglion  so  definite  that  its  confusion  with 
all  other  cell  groups  will  be  impossible,  and  if  this  result  be  obtained 
there  need  be  little  concern  as  to  the  fitness  of  the  name  employed. 

Location  and  extent  of  the  ganglion  opticuni  basale. 

MAX. 

The  basal  optic  ganglion  in  man  is  a  mass  of  gray  matter,  super- 
ficially situated  partly  in  the  tuber  cinereum  and  partly  in  the  anterior 
perforated  substance,  which  extends  along  both  borders  of  the  optic 
tract.  The  main  mass  of  cells  forms  in  the  anterior  j>erforated 
substance  a  column  which  is  closely  applied  to  the  dorsal  portion  of  the 
oro-lateral  surface  of  the  optic  tract.  This  mass  is  connected  by  a 
comparatively  small  number  of  scattered  cells  situated  directly  dorsal 
to  the  tract  with  another  cell  mass  which  follows  in  the  tuber  cinereum 
the  caudo-medial  border  of  the  optic  tract ;  in  other  words  the  basal  optic 
ganglion  consists,  generally  speaking,  of  two  parallel  columns  of  cells 
lying  along  either  side  of  the  optic  tract,  which  are  connected  by 
scattered  cells  of  the  same  type  lying  dorsal  to  the  optic  tract.  For  the 
second  time  I  strongly  recommend  a  consideration  of  pp.  39-40,  where 
it  is  pointed  out  just  what  each  type  of  illustration  is  intended  to  show. 

Taking  up  the  first  series  of  man  (Series  AC)  the  basal  optic  gang- 
lion appears  caudally  in  Fig.  4  as  a  few  scattered  cells  situated  on  the 
peripher}-  of  the  tuber  cinereum ;  Figs.  5  and  6  show  the  development 
of  the  cell  mass  as  we  pass  orally.  The  cell  masses  shown  in  Figs.  4 
and  5  are  to  be  regarded  as  a  caudal  projection  of  the  cell  column  lying 
along  the  caudo-medial  border  of  the  optic  tract,  and  are  continuous 
with  the  caudal  portion  of  this  cell  column  which  appears  first  in  Fig.  6. 
Between  Figs.  6  and  7  is  a  considerable  gap,  in  which  the  following 


Xuclci  Tiiberis  Latcrales  and  the  Ganglion  Opticum  Basale.       9 

changes  have  occurred :  as  the  optic  tract  approaches  the  median  plane 
and  applies  itself  to  the  base  of  the  brain  the  cell  mass  continues  to 
increase  in  size,  extending  further  laterally  and  also  to  a  less  extent 
further  medially,  and  assumes  a  position  as  a  column  on  the  dorsal 
surface  of  the  caudo-medial  aspect  of  the  optic  tract,  or  in  other  words 
it  extends  in  the  tuber  cinereum  along  the  line  where  the  tuber  joins 
the  optic  tract.  Midway  between  Figs.  6  and  7  the  medial  pole  of  the 
column  crosses  the  median  line  and  caudal  to  the  optic  chiasm  unites 
with  the  corresponding  column  of  the  other  side  by  means,  however,  of 
only  a  few  scattered  cells  situated  in  the  iirfundibulum.  The  next 
series  (D)  shows  this  fact.  But  still  another  change  has  occurred 
between  Figs.  6  and  7,  namely,  after  reaching  its  greatest  development, 
which,  as  previously  described,  occurs  at  the  juncture  of  the  tuber 
cinereum  with  the  caudo-medial  border  of  the  optic  tract,  the  cell  mass 
becomes  rapidly  diminished  to-  only  a  few  scattered  cells  which  are 
situated  on  the  dorsal  (or  deep)  surface  of  the  optic  tract.  Fig.  7  shows 
the  extent  of  the  basal  optic  ganglion  slightly  oral  to  this  region  of  its 
poorest  development;  the  section  passes  through  the  optic  tract  near 
its  oro-lateral  border  and  since  the  section  is  near  this  border  the  cell 
mass  has  increased  in  extent.  Between  Figs.  7  and  8  it  increases 
steadily  in  size  to  reach  its  maximum  in  Fig.  8 ;  this  figure  represents 
a  section  through  the  cell  column  which  extends  in  the  anterior  perfor- 
ated substance  along  the  oro-lateral  border  of  the  optic  tract.  In  Fig.  9 
the  basal  optic  ganglion  has  diminished  in  size,  and  this  is  due  to  a 
shortening  of  the  lateral  portion  of  the  column ;  that  the  lateral  portion 
of  the  column  should  be  shortened  is  evident  when  we  recall  that  the 
cell  column  lies  parallel  to  the  optic  tract,  and  that  the  course  of  the 
optic  tract  is  such  that  in  the  present  plane  of  section  as  one  proceeds 
orally  the  lateral  portion  of  the  tract  appears  and  disappears  first  (see 
Figs.  1  to  10).  Fig.  10  shows  the  most  oral  portion  of  the  basal  optic 
ganglion.  The  fact  that  the  portion  of  the  ganglion  situated  caudal 
(medial)  to  the  optic  tract  does  not  extend  as  far  laterally  -as  that  por- 
tion situated  oral  (lateral)  to  the  tract  should  be  correlated  with  the 
difference  of  the  relation  of  the  two  borders  of  the  tract  to  the  base  of 
the  brain ;  for  while  the  oral  (lateral)  border  of  the  tract  is  in  intimate 
relation  to  the  anterior  perforated  substance  from  the  chiasm  to  a  point 
far  lateral  from  the  median  plane,  the  caudal  (medial)  border  of  the 
tract  is  in  intimate  relation  to  the  tuber  cinereum,  which  extends  for 


10  Edward  F.  M alone. 

a  shorter  distance  laterally,  and  then  the  tract  passes  over  the  cms 
cerebri,  where  of  course  it  is  no  longer  in  relation  to  gray  matter. 

To  sum  up.  Series  AC  of  man  (Figs.  4  to  10)  shows,  in  sections 
almost  parallel  to  the  course- of  the  optic  tract,  the  appearance  of  the 
three  parallel  columns  of  cells  which  constitute  the  basal  optic  gang- 
lion ;  one  column  lies  in  the  tuber  cinereum  along  the  medio-caudal 
border  of  the  optic  tract,  the  second  extends  along  the  dorsal  (deep) 
surface  of  the  tract  and  consists  of  only  a  few  cells,  while  the  third  and 
largest  column  extends  in  the  anterior  perforated  substance  along  the 
latero-oral  border  of  the  tract.  The  caudal  column  is  continuous  in  the 
infundibulum  with  the  corresponding  column  of  the  opposite  side.  The 
figures  show  the  inevitable  relation  of  three  parallel  columns  in  slightly 
oblique  section. 

Series  D  of  man  (Figs.  11  to  18)  shows  essentially  the  same  relations, 
but  the  plane  of  section  is  different.  In  the  previous  series  the  plane 
was  almost  parallel  to  the  course  of  the  optic  tract,  although  slightly 
approaching  the  plane  of  a  cross  section ;  in  Series  D  the  plane  of  sec- 
tion is  almost  at  right  angles  to  the  course  of  the  optic  tract,  and  both 
the  caudo-medial  and  the  oro-lateral  borders  are  shown  in  all  sections. 
It  is  evident  that  a  plane  of  section  at  right  angles  to  the  median  plane 
(which  would  cut  both  halves  of  the  brain  symmetrically)  could  not 
pass  through  the  optic  tract  at  right  angles  to  its  course,  since  the  two 
tracts  converge  towards  the  median  plane  as  they  pass  orally ;  therefore 
to  obtain  a  cross  section  of  the  right  tract  the  plane  of  section  must  lie 
more  oral  on  this  side  than  on  the  left.  The  asymmetry  of  the  plane 
of  section  of  Series  D  is  not  quite  great  enough  to  produce  a  cross  sec- 
tion of  the  right  optic  tract,  although  this  condition  is  very  nearly 
attained. 

When  Fig.  11  of  Series  D  is  compared  with  Fig.  4  of  Series  AC  the 
location  of  the  basal  optic  ganglion  is  rather  confusing,  for  in  the 
latter  series  it  appeared  oaudally  in  the  tuber  cinereum  near  the  medio- 
caudal  border  of  the  optic  tract,  while  in  Fig.  11  it  appears  first  in  the 
anterior  perforated  substance  along  the  oro-lateral  border;  if  we  keep 
in  mind  the  asymmetry  of  the  section,  and  the  fact  that  the  cell  column 
here  shown  extends  further  laterally  than  the  other  two  columns  of  the 
cell  group  and  that  in  such  asymmetrical  sections  the  lateral  portion  is 
the  most  oral,  little  difficulty  should  exist  in  realizing  just  why  this 
one  of  the  three  cell  columns  should  appear  first.  Fig.  12  shows  practi- 
cally the  game  relations,  except  that  the  optic  tract  is  nearer  the  median 


Nuclei  Tuheris  Laterales  and  the  Ganglion  Opticum  Basale.     11 

line.  In  Fig.  13  the  cell  column  along  the  medic-caudal  border  of  the 
tract,  as  well  as  the  scattered  cells  dorsal  to  the  tract,  appears,  and 
from  here  throughout  the  entire  series  these  three  columns  are  present. 
Of  course  all  columns  are  in  cross  section.  In  Fig.  18  the  cell  mass  is 
seen  to  extend  across  the  median  line,  a  condition  which  occurs  also  in 
the  three  other  series  of  man.  Series  D  (Figs.  11  to  18)  shows  clearly 
that  the  basal  optic  ganglion  consists  of  two  well  developed  parallel 
columns  of  cells  connected  by  an  intermediate  parallel  column  consisting 
of  only  a  few  cells.  As  will  be  shown  later  the  exact  extent  of  the 
basal  optic  ganglion  in  all  the  mammals  studied  is  readily  determined 
by  the  distinctive  histological  character  of  its  cells,  and  accordingly  the 
three  parallel  cell  columns  have  been  considered  as  merely  subdivisions 
of  one  nucleus,  not  because  this  seems  convenient,  but  because  the 
identical  character  of  their  cells  makes  such  a  conclusion  unavoidable. 

MAC.A.CUS  RHESUS. 

In  maeaeus  rhesus  the  basal  optic  ganglion  is  shown  first  in  Fig.  22, 
although  it  extends  slightly  further  caudally.  In  tliis  figure  the  gang- 
lion is  situated  just  oral  to  the  line  of  apposition  between  the  tuber 
cinereum  and  the  optic  tract.  In  the  sections  between  Figs.  23  and  23, 
as  we  pass  orally,  the  cells  become  reduced  in  number  and  lie  dorsal  to 
the  tract,  and  as  the  oro-lateral  border  of  the  tract  is  reached  increase 
in  number.  In  Fig.  23  appears  the  cell  column  which  lies  in  the  ante- 
rior perforated  substance  along  the  oro-lateral  border  of  the  tract.  The 
further  development  of  the  ganglion  (Figs.  21:  and  25)  needs  no 
description.  In  Fig.  22  the  medial  pole  of  the  ganglion  is  seen  to 
approach  the  median  line,  and  between  Figs.  22  and  23,  just  caudal  to 
the  optic  chiasm,  a  very  few  widely  scattered  cells  unite  the  ganglia  of 
opposite  sides.  As  in  man  the  basal  optic  ganglion  in  macacus  is 
seen  to  consist  of  tlie  same  three  parallel  columns  of  cells,  of  which  the 
oro-lateral  column  (situated  in  the  anterior  perforated  substance)  con- 
stitutes the  greater  part  of  the  ganglion. 

LEMDR   RUFUS. 

The  basal  optic  ganglion  of  the  lemur  does  not  differ  essentially 
from  that  of  man  and  maoacus,  except  that  whereas  in  the  two  latter 
animals  it  consisted  of  two  parallel  cell  columns  loosely  connected  by 
a  third,  in  the  lemur  this  connecting  mass  of  scattered  cells  (dorsal  to 


12  Edward  F.  M alone. 

the  optic  tract)  is  missing;  consequently  the  ganglion  consists  of  two 
entirely  separate  parallel  columns  of  cells.  Fig.  27  represents  a  section 
between  the  two  cell  columns.  As  in  man  and  macacus  the  ganglia  of 
opposite  sides  are  united  by  a  very  few  cells;  this  union  occurs  just 
caudal  to  the  optic  chiasm  (slightly  caudal  to  the  level  represented 
in  Fig.  27). 

CAT. 

The  basal  optic  ganglion  in  the  cat  (Figs.  31  to  35)  has  practically 
the  same  location  and  extent  as  in  the  lemur.  As  in  the  lemur  the  gang- 
lion consists  of  the  same  two  parallel  cell  columns,  which  are  completely 
separate.  Between  Figs.  32  and  33  lies  the  region  in  which  the  gang- 
lion is  absent.  Just  caudal  to  the  optic  chiasm  (caudal  to  level  of  Fig. 
33)  the  ganglia  of  opposite  sides  are  united  in  this  series  by  a  few 
cells ;  in  the  other  two  series  of  the  cat  no  union  was  present. 

Comparing  the  location  and  extent  of  the  basal  optic  ganglion  in  all 
four  animals  the  following  facts  should  be  noted: 

1.  The  basal  optic  ganglion  in  all  four  animals  consists  almost 
exclusively  (in  some  cases  exclusively)  of  two  compact,  parallel  cell 
columns.  The  larger  of  these  two  columns  lies  superficially  in  the 
anterior  perforated  substance  along  the  line  where  this  region  becomes 
continuous  with  the  oro-lateral  border  of  the  optic  tract ;  the  smaller 
column  lies  superficially  in  the  tuber  cinereum  along  the  line  where  it 
joins  the  optic  tract. 

2.  These  two  constant,  parallel  cell  columns  are  united  in  man,  and 
to  a  less  extent  also  in  macacus,  by  more  or  less  diffusely  scattered  cells 
of  the  same  type  which  lie  dorsal  to  the  optic  tract. 

3.  In  all  four  forms  there  occurs  a  union  of  the  ganglia  of  opposite 
sides  by  means  of  diffusely  scattered  cells  located  just  caudal  to  the 
optic  chiasm.  In  man  this  union  is  very  definite,  in  macacus  less 
definite,  while  in  the  other  two  animals  it  is  rudimentary  and  due  to  the 
presence  of  a  very  few  widely  scattered  cells  between  the  two  ganglia. 
In  the  cat  even  this  feebly  developed  fusion  is  not  present  in  all 
individuals. 

4.  Although  the  phylogenetic  series  from  man  to  the  cat  is  too  short 
to  be  of  much  service  in  revealing  the  phylogenetic  development  of  so 
constant  a  cell  group  as  the  basal  optic  ganglion,  it  is  apparent  that 
as  we  descend  the  series  the  two  parallel  cell  columns  become  separate 
(more  closely  united  in  man  than  in  macacus,  and  separate  in  the  lemur 


Nuclei  Tuberis  Laterales  and  the  Ganglion  Opticum  Basale.     13 

and  cat)  ;  the  same  is  trae  as  to  the  fusion  of  the  ganglia  of  opposite 
sides  (definite  in  man,  less  so  in  macacus,  barely  present  in  the  lemur 
ajid  not  always  present  in  the  cat.) 

Cell  type  of  the  basal  optic  ganglion. 

It  is  not  my  intention  to  attempt  to  give  in  words  the  characteristics 
of  the  cells  of  the  basal  optic  ganglion  which  have  been  already  satis- 
factorily shown  in  the  illustrations.  I  shall  confine  myself  to  pointing 
out  the  fundamental  characters  of  these  cells,  and  when  we  have  thus 
become  familiar  with  the  main  features  of  the  cell  picture  we  shall  be 
in  a  position  to  discuss  the  relations  of  these  cells  to  those  of  other 
groups.  The  water  color  reproductions  of  the  cells  of  the  basal  optic 
ganglion,  and  of  all  other  cells  thus  illustrated  (Figs.  38  to  58),  were 
all  drawn  from  cross-sections  of  the  brain.  In  Series  D  of  man,  in 
which  the  plane  of  section  differs  widely  from  that  of  a  cross-section, 
I  have  been  unable  to  observe  any  important  difference  in  the  appear- 
ance of  the  cells  of  the  basal  optic  ganglion,  and  it  is  my  opinion  that 
the  plane  of  section  does  not  affect  to  any  appreciable  extent  the  appear- 
ance of  the  majority  of  these  cells.  The  cells  of  the  basal  optic  gang- 
lion and  of  all  other  groups  illustrated  are  as  characteristic  in  Series  D 
a.s  in  Series  AC.  However,  I  have  not  made  a  careful  enough  study 
of  this  point  to  state  definitely  that  the  cell  character  is  absolutely 
unaffected  by  differences  in  the  plane  of  section.  The  cells  of  the  basal 
optic  ganglion  in  all  four  animal  forms  (Figs.  38,  44,  50  and  55)  are 
large  polygonal  cells  which  possess  very  coarse  processes.  These  pro- 
cesses, as  is  shown  in  Fig.  37,  form  an  intercellular  feltwork;  under 
low  power  one  would  hardly  suppose  that  this  feltwork  was  composed 
of  cell  processes,  and  the  ganglion  appears  to  be  characterized  not  only 
by  its  typical  cells  but  also  by  the  presence  of  a  distinctive  intercellular 
substance.  These  processes  are  practically  colorless,  and  are  in  Fig. 
37  represented  as  blue  because  at  this  magnification  they  could  not  be 
shown  in  any  other  manner.  The  cell  processes  in  Figs.  38,  44,  50  and 
55  are  not  as  long  as  they  appear  in  the  actual  preparations.  Another 
fundajnental  character  of  these  cells  is  the  distribution  of  the  Nissl 
substance,  nearly  all  of  which  is  massed  on  the  periphery  of  the  cell ; 
this  peripheral  distribution  is  not  equal  in  all  portions  of  the  periphery, 
since  the  depth  is  much  greater  at  certain  points,  and  at  other  points 
of  the  periphery  the  iNrissl  substance  may  be  almost  entirely  absent. 


14  ,  Edicard  F.  Malone. 

Of  course  the  distribution  of  the  Xissl  substance  will  appear  different 
according  to  the  location  of  the  optical  section,  and  the  figures  show  this 
to  some  extent,  although  they  represent  to  a  certain  extent  a  combination 
of  different  optical  sections.  Although  minor  differences  in  cell  char- 
acter exist  in  different  animals,  the  cell  group  is  phylogenetically 
so  old  that  in  the  relatively  brief  interval  between  man  and  the  cat  no 
changes  in  cell  character  have  occurred  which  are  sufficiently  funda- 
mental to  permit  of  correlation  with  the  phylogenetic  position  of  the 
corresponding  animals ;  this  point  will  be  discussed  later  in  connection 
with  the  nuclei  tuberis,  where  such  a  relation  between  cell  character 
and  the  phylogenetic  position  of  tlie  corresponding  animal  actually 
exists.  The  structure  of  the  cells  of  the  ganglion  opticum  basale  is 
such  as  to  exclude  the  possibility  of  these  cells  being  motor,  whereas 
the  large  size  of  the  cells  probably  indicates  (as  Dr.  Donaldson  has 
suggested  to  me)  that  they  either  receive  impulses  converging  from 
many  sources  or  distribute  impulses  over  an  extensive  region. 

Separation  of  the  hasal  optic  ganglion  from  surrounding  cell  groups 
through  differences  in  cell  ch^aracier. 

A  subdivision  of  any  portion  of  the  nen'ous  system  based  merely 
upon  the  splitting  up  of  gray  matter  through  the  mechanical  agency  of 
fiber  masses  is,  except  in  certain  cases,  valuable  merely  with  reference 
to  orientation;  such  a  subdivision  should  therefore  be  considered  as  a 
crude  (although  necessary)  beginning  to  be  followed  by  a  more  careful 
study  of  the  region  involved  in  which  the  cell  character  of  the  various 
groups  receives  careful  attention.  Moreover  it  is  highly  unsatisfactory 
for  an  author  to  state  dogmatically  that  he  recognizes  the  presence  of 
certain  cell  groups  upon  the  basis  of  certain  differences  of  cell  char- 
acter, concerning  which  he  is  either  silent  or  else  treats  in  a  superficial 
manner;  for  we  are  left  in  doubt  as  to  whether  these  differences  in  cell 
character  are  really  fundamental,  and  as  to  the  degrees  of  relationship 
between  the  cell  characters  of  various  cell  groups.  When  we  have  de- 
scribed the  location  and  extent  of  various  cell  groups  and  have  made  it 
possible  to  recognize  this  by  means  of  a  definite  cell  type,  and  when  we 
have  clearly  shown  the  differences  and  similarities  in  cell  character  of 
various  cell  groups,  and  have  pointed  out  the  phylogenetic  development 
and  relations  of  these  groups,  then  and  not  until  then  will  there  be  a 
basis  for  experimental  work  which  will  help  solve  many  important 


Nuclei  Tuheris  Laterales  and  the  Ganglion  Opticum  Basal e.     15 

questions  as  to  the  elementary  mechanisms  of  the  nervous  system.  In 
the  case  of  the  basal  optic  ganglion,  therefore,  I  shall  not  be  content 
with  having  pointed  out  its  location  and  extent,  but  shall  proceed  to 
compare  the  character  of  its  cells  with  that  of  the  cells  of  surrounding 
groups ;  such  a  comparison  serves  not  only  to  set  aside  this  cell  group 
as  different  from  other  cell  groups  by  virtue  of  differences  in  cell 
character,  but  it  enables  us  also  to  recognize  certain  similarities  of  cell 
character  between  the  cells  of  the  ganglion  and  the  cells  of  other  groups, 
so  that  different  degrees  of  relationship  between  the  different  cell 
groups  may  be  provisionally  stated. 

I  shall  first  point  out  the  differences  in  cell  character  between  the 
cells  of  the  basal  optic  ganglion  and  those  of  the  surrounding  groups 
so  .as  to  complete  the  picture  of  this  cell  group ;  afterwards  the  relations 
of  various  cell  groups  will  be  discussed  (but  not  until  the  nuclei 
tuberis  have  been  described).  There  is  no  diiSculty  in  distinguishing 
the  cells  of  the  basal  optic  ganglion  from  those  of  the  nuclei  tuberis 
and  of  the  substantia  grisea  ventriculi  tertii.  This  is  shown  by  a 
reference  to  the  corresponding  figures  38,  39  and  -iO;  4-1,  45  and  46;  50 
and  51;  55  and  56;  moreover  the  difference  between  the  cells  of  the 
basal  optic  ganglion  and  those  of  the  substantia  grisea  is  well  shown  in 
Fig.  37. 

The  cells  of  the  basal  optic  ganglion  may  be  readily  distinguished 
also  from  those  of  the  nucleus  tubero-mammillaris,  although  both  types 
of  cells  have  a  certain  similarity;  the  failure  to  recognize  the  unity  of 
the  cell  complex  known  as  the  nucleus  tubero-mammillaris  (mammillo- 
infundibularis)  and  to  clearly  separate  it  from  the  basal  optic  ganglion 
is  one  of  the  most  important  causes  for  our  meager  knowledge  of  this 
region.  Considering  first  the  relations  of  these  tvi'o  cell  groups  as  to 
location,  it  is  evident  from  the  illustrations  that  they  are  closely  related 
only  for  a  short  distance.  Moreover  the  cells  of  the  nucleus  tubero- 
jnammillaris  are  for  the  most  part  rather  diffusely  scattered,  while 
those  of  the  basal  optic  ganglion  are  densely  packed  together  and  the 
cell  group  is  further  characterized  by  the  intercellular  feltwork  formed 
by  the  coarse  colorless  cell  processes ;  these  two  points  may  be  observed 
by  comparing  Figs.  36  and  37.  Comparing  the  cell  type  of  the  two  cell 
groups  the  fundamental  difference  found  in  all  four  animals  involves 
the  appearance  of  the  Nissl  substance ;  in  both  forms  the  Nissl  substance 
is  located  principally  on  the  periphery  of  the  cell,  but  in  the  cells  of  the 
basal  optic  ganglion  the  massing  of  the  Nissl  substance  on  the  periphery 

2 


16  Edward  F.  Malone. 

is  extreme,  whereas  in  the  cells  of  the  nucleus  tubero-mammillaris  the 
N"issl  substance  is  not  so  densely  packed  together  on  the  periphery  and 
more  of  it  is  present  in  the  central  portion  of  the  cell.  In  other  words 
the  Nissl  substance  in  the  cells  of  the  nucleus  tubero-mammillaris  is 
more  diffusely  distributed  throughout  the  entire  cell.  The  difference 
in  cell  type  obtains  throughout  the  entire  extent  of  both  nuclei.  A 
study  of  the  illustrations  shows  that  while  other  differences  in  cell  type 
occur  in  certain  animals  the  fundamental  difference  consists  in  the 
mode  of  distribution  of  the  Nissl  substance  (Figs.  38  and  41 ;  44  and 
47;  50  and  52;  55  and  57). 

The  separation  of  the  basal  optic  ganglion  from  the  nucleus  paraven- 
tricularis  hypothalami  on  the  basis  of  cell  type  is  difficult.  Fortunately 
both  are  in  all  four  animals  sharply  circumscribed,  and  although  the 
extremities  of  these  two  cell  columns  approach  one  another  (Fig.  10), 
they  never  actually  fuse.  A  study  of  the  location  of  these  two  cell 
groups  will  show  their  axes  are  almost  at  right  angles  to  each  other. 
In  a  former  article  I  made  the  statement  that  the  cells  of  the  basal 
optic  ganglion  in  man  could  not  be  distinguished  from  those  of  the 
nucleus  paraventricularis,  but  a  more  careful  study  of  the  cell  groups 
in  man  together  with  their  study  in  other  animals  proves  that  this 
statement  is  not  correct,  although  not  far  from  the  truth.  As  a  matter 
of  fact  I  have  been  unable  to  observe  any  one  fundamental  difference 
between  these  two  types  of  cells  which  is  clearly  shown  in  all  four 
animal  forms,  although  in  each  animal  some  differences  are  present 
which  always  make  a  distinction  possible.  There  is  one  difference  which 
holds  fairly  well  for  all  forms,  and  this  is  the  same  difference  that  was 
so  evident  between  the  cells  of  the  basal  optic  ganglion  and  those  of  the 
nucleus  tubero-mammillaris,  namely,  the  Xissl  substance  is  more 
densely  massed  on  the  periphery  of  the  cell  in  the  case  of  the  basal  optic 
ganglion,  while  in  the  cells  of  the  nucleus  paraventricularis  the  Nissl 
substance  is  more  uniformly  distributed  throughout  the  cell.  In  man 
( Figs.  38  and  43 )  this  difference  is  not  marked,  but  in  the  basal  optic 
ganglion  (Fig.  38)  the  Xissl  substance  on  the  ]3eriphery  forms  in  part 
large,  irregularly  placed  masses  of  granules,  while  in  the  cells  of  the 
nucleus  paraventricularis  (Fig.  43)  the  granules  are  smaller,  more 
uniform  in  size  and  do  not  stain  so  intensely;  at  the  same  time  the 
c-entral  portion  of  the  cell  is  somewhat  more  deeply  stained  (due  to  more 
Nissl  substance)  than  in  the  case  of  the  cells  of  the  basal  optic  ganglion. 
Other  differences  in  man  are  as  follows:  the  cells  of  the  basal  optic 


Nuclei  Tuberis  Laterales  and  the  Ganglion  Opticum  Basale.     17 

ganglion  are  somewhat  larger  (the  smallest  cell  illustrated  in  Pig.  38 
shows  only  a  small  portion  of  a  cell  in  optical  section),  the  nuclear 
membrane  is  not  so  definite,  and  the  nucleus  is  larger.  In  macacus 
(Figs.  44  and  49)  the  difference  in  distribution  of  the  Nissl  substance 
is  less  than  in  any  of  the  other  animals,  but  even  here  the  Nissl  sub- 
stance in  the  cells  of  the  l>asal  optic  ganglion  is  more  nearly  confined 
to  the  periphery,  is  denser  here  and  not  so  dense  in  the  interior  portion 
of  the  cell  as  in  the  case  of  the  other  cell  group;  this  results  in  a 
sharper  differentiation  between  peripheral  and  central  portions  of  the 
cell  in  the  case  of  the  ba&al  optic  ganglion.  Other  differences  in 
macacus  are:  the  cells  and  cell  nuclei  of  the  basal  optic  ganglion  are 
smaller  than  those  of  the  nucleus  paraventricularis  (the  opposite  is 
true  in  man),  and  the  cells  of  this  latter  nucleus  are  rather  piriform. 
In  the  lemur  (Figs.  50  and  54)  the  difference  in  distribution  of  the 
Nissl  substance  is  marked,  the  differentiation  of  the  cell  into  peripheral 
and  central  portions  being  much  sharper  in  the  cells  of  the  basal  optic 
ganglion.  In  the  cat  (Figs.  55  and  58)  the  difference  between  per- 
iphery and  center  of  the  cell  is  again  more  marked  in  the  cells  of  the 
basal  optic  ganglion,  since  in  the  cells  of  the  other  group  the  central 
portion  of  the  cell  (including  the  cell  nucleus)  stains  deeply  (Fig.  58). 
Note  also  in  the  cat  that,  just  as  in  man,  the  cells  of  the  basal  optic 
ganglion  are  larger,  but  unlike  those  of  man  have  smaller  cell  nuclei 
than  the  cells  of  the  nucleus  paraventricularis. 

The  resemblance  of  the  cell  type  in  the  case  of  the  basal  optic  gang- 
lion and  the  nucleus  paraventricularis  hypothalami  is  therefore  very 
close,  but  in  all  four  animals  studied  the  cells  of  the  basal  optic  gang- 
lion were  found  to  be  more  sharply  differentiated  into  a  peripheral  zone 
containing  dense  masses  of  Nissl  substance  and  an  inner  portion  rela- 
tively free  from  this  substance,  whereas  in  the  cells  of  the  nucleus  par- 
aventricularis hypothalami  the  Nissl  substance  was  not  so  sharply 
differentiated  as  to  its  distribution.  It  will  be  recalled  that  within  the 
cells  of  the  nucleus  tubero-mammillaris  this  differentiation  was  even 
less  marked  than  within  those  of  the  nucleus  paraventricularis. 

Another  cell  group  must  be  distinguished  from  the  basal  optic  gang- 
lion. This  is  the  nucleus  ansae  peduncularis  of  Meynert  (ganglion 
basale  of  Kolliker) .  It  occurs  in  man,  macacus  and  the  lemur,  but  I 
was  unable  to  find  it  in  any  of  my  three  series  of  the  cat.  Its  location 
and  extent  may  be  seen  in  the  different  illustrations.  In  all  three 
animals  in  which  it  occurs  it  is  readily  distinguished  from  the  basal 


18  Edward  F.  Malone. 

optic  ganglion  by  tlie  large  size  of  its  cells.  In  man,  as  was  pointed 
out  by  Kolliker,  the  cells  of  the  ganglion  basale  may  be  readily  separated 
from  those  of  the  basal  optic  ganglion  by  the  fact  that  they  are  heavily 
pigmented  (Fig.  42).  In  a  previous  article  I  have  discussed  the  value 
of  pigmentation  as  a  basis  of  distinguishing  different  cell  types. 
Yellow  pigment  occurs  in  certain  nerve  cells  of  the  adult  human  and 
increases  with  age,  but  this  occurrence  and  increase  in  amount  is  not 
errntic,  but  behaves  differently  with  respect  to  different  types  of  cells. 
Some  types  of  cells  never  contain  pigment,  others  always  contain  it, 
while  still  other  types  may  contain  none  or  under  other  conditions  may 
contain  a  small  or  even  a  moderate  amount.  But  however  the  total 
amount  of  yellow  pigment  in  the  brain  of  different  adults  may  vary, 
the  amount  in  each  specific  type  of  cell  retains  the  same  relation  to  that 
of  every  other  type  of  cell ;  in  other  words,  the  relative  amount  of  pig- 
ment in  the  different  types  of  cells  is  constant  in  all  individuals.  Note 
that  this  pigmentation  of  the  cells  of  the  ganglion  basale  in  man  is 
accompanied  by  an  additional  characteristic  (large  size)  and  that  the 
homologous  cells  of  macacus  and  the  lemur  are  entirely  different  from 
those  of  other  cell  groups,  although  pigmentation  is  of  course  lacking. 
This  question  of  the  relative  amount  of  yellow  pigment  as  a  character- 
istic of  certain  types  of  cells  which  differ  also  in  other  respects  will  be 
noted  again  in  connection  with  the  nuclei  tuberis.  This  yellow  pigment 
should  not  be  confounded  with  the  brown  pigment  which  occurs  in  the 
cells  of  the  substantia  nigra  and  elsewhere. 

Now  that  the  differences  in  cell  type  between  the  cells  of  the  basal 
optic  ganglion  and  those  of  surrounding  groups  have  been  studied  it 
woiild  be  desirable  to  consider  the  relationship  of  these  various  cell 
groups  from  the  standpoint  of  similarities  in  cell  character.  This 
consideration,  as  well  as  that  of  the  literature,  must  however  be  post- 
poned until  the  nuclei  tuberis  have  been  described,  since  such  a  com- 
parison of  the  various  cell  groups  demands  a  familiarity  with  the 
nuclei  tuberis  and  the  substantia  grisea  of  the  third  ventricle. 

Nuclei  Tuberis  Later-^les. 

Under  the  name  of  nuclei  tuberis  laterales  I  shall  describe  certain 
cell  groups  of  the  pars  optica  hypothalami  (telenceplialon)  which  I 
have  formerly  considered  very  briefly  in  my  monograph  on  the  human 
dieneephalon.     Although  a  number  of  authors  have  written  of  nuclei 


Nuclei  Tuheris  Laterales  and  the  Ganglion  Opticuin  Basale.     19 

tuberis,  except  in  one  case,  there  is  absolutely  no  reason  to  believe  that 
any  of  them  have  had  in  mind  the  characteristic  nests  of  cells  which 
will  now  be  described.  Kolliker  undoubtedly  saw  these  nuclei, 
although,  as  will  appear  later,  his  description  is  faulty  and  his  figures 
incorrectly  labeled ;  the  result  is  so  confusing  that  a  careful  study  of 
this  region  is  necessaiy  to  appreciate  the  value  of  Kolliker's  observa- 
tions. In  the  previous  brief  description  I  employed  the  name  nuclei 
tuberis,  but  in  the  present  article  adopt  the  name  nuclei  tuberis  later- 
ales.  The  name  nuclei  tuberis  is  nor  distinctive  and  might  be  used  to 
indicate  any  cell  group  whatsoever  located  in  the  tuber  cinereum;  as 
a  matter  of  fact  it  has  been  so  used,  and  the  resulting  confusion  may  be 
imagined  when  one  considers  how  many  different  cell  groups  lie  in  the 
tuber  cinereum.  This  confusion  is  not  one  merely  of  names,  but 
involves  the  failure  to  distinguish  (regardless  of  names)  various  cell 
groups  of  the  tuber  cinereum  which  in  properly  prepared  material  may 
be  readily  distinguished  not  only  through  differences  in  location  but 
also  through  striking  differences  in  cell  type. 

Location  and  extent  of  the  nuclei  tuberis  laierales. 

The  nuclei  tuberis  laterales  consist  in  man  and  macacus  of  several 
nests  of  cells  located  on  the  periphery  of  the  tuber  cinereum.  In  the 
lemur  in  this  location  one  sees  the  beginning  differentiation  of  these 
cell  groups  from  the  cells  which  surround  them,  although  these  differ- 
ences are  so  slight  that  I  have  not  attempted  to  show  them  in  the  illus- 
trations.   In  the  cat  these  cell  groups  are  absent. 

MAN. 

In  Series  AC  of  man  the  nuclei  tuberis  laterales  are  shown  in  Figs. 
1  to  8;  the  size  of  the  cells,  however,  has  been  so  much  reduced  that 
they  should  be  examined  with  the  aid  of  a  hand  lens.  In  Pig.  1  two 
indentations  are  shown  on  the  base  of  the  brain  which  should  be 
noted.  One  indentation  marks  off  on  the  base  of  the  brain  the  medial 
boundary  of  the  pes  pedunculi,  while  the  more  medial  indentation 
forms  the  superficial  boundary  between  the  tuber  cinereum  and  the 
nucleus  tubero-mammillaris  (nucleus  mammillo-infundibularis),  this 
nucleus  lying  between  these  two  fissures.  The  area  bounded  by  these 
two  fissures  ventrally,  and  by  the  columna  fornicis  dorsally,  is  of 
especial  importance  in  describing  the  location  of  the  nuclei  tuberis. 


20  Edward  F.  Malone. 

On  examining  Figs.  1  to  6  these  two  fissures  are  seen  to  be  practically 
parallel,  and  the  area  bounded  by  them  and  by  the  fornix  column  under- 
goes as  we  pass  orally  the  following  change :  in  the  caudal  portion  (Fig. 
1 )  this  area  is  occupied  by  the  cells  of  the  nucleus  tubero-mammillaris, 
whose  caudal  pole  constitutes  the  lateral  ganglion  of  the  maniraillary 
body;  passing  further  orally  these  large  cells  are  replaced  (especially 
those  situated  ventrally  and  medially)  by  the  small  cells  of  the  nuclei 
tuberis  laterales  and  of  the  substantia  grisea.  Accordingly  the  oaudal 
portion  of  this  region  is  continuous  with  the  lateral  ganglion  of  the 
mammillary  body,  whereas  orally  it  gradually  becomes  the  lateral 
portion  of  the  tuber  cinereum.  In  this  region  included  between  these 
two  fissures  ventrally,  the  pes  pedunculi  laterally,  and  the  fornix 
column  dorsally,  lies  by  far  the  greater  portion  of  the  cells  of  the  nuclei 
tuberis  laterales.  Referring  to  Figs.  1  to  8  the  location  and  extent  of 
these  cell  groups  are  evident,  and  only  a  few  comments  will  be  necessary. 
The  nuclei  tuberis  laterales  consist  of  a  variable  number  of  well  circum- 
scribed nests  of  small  cells,  and  can  be  distinguished  even  without  the 
aid  of  their  characteristic  cell  type.  These  different  cell  nests  are 
more  or  less  connected  with  one  another,  and  their  separation  is  to  be 
regarded  probably  as  dependent  upon  the  mechanical  influence  of  fiber 
masses,  since  their  partial  fusion,  identical  cell  type,  -and  variable 
number  would  seem  to  exclude  the  possibility  of  these  separate  groups 
having  different  functions.  The  cell  nests  of  the  nuclei  tuberis  later- 
ales are  practically  free  from  cells  of  surrounding  groups. 

In  considering  the  second  series  of  man  (Series  D,  Figs.  11  to  15)  it 
is  absolutely  essential  to  understand  the  plane  of  sectionr;  this  has 
already  been  described  in  considering  the  basal  optic  ganglion.  It  will 
suffice  to  point  out  that  Series  D  differs  from  Series  AC  in  that  the 
plane  of  section  of  the  former  passes  more  caudally  as  it  passes  from 
dorsal  to  ventral ;  and  that  in  it  the  opposite  sides  of  the  brain  are  cut 
asymmetrically,  in  that  the  lateral  portion  of  each  section  of  Series  D 
is  situated  further  oral  than  the  medial  portion  of  the  corresponding 
section.  As  was  previously  noted  the  plane  of  section  of  Scries  D  is  the 
only  one  which  can  pass  through  the  right  optic  tract  at  right  angles 
to  its  long  'axis ;  this  relation  to  the  optic  tract  will  render  a  clear  picture 
of  the  plane  of  section  relatively  easy.  Bearing  this  plane  o^  section 
in  mind  a  study  (with  the  aid  of  a  hand  lens)  of  Figs.  11  to  15  will 
show  that  the  location  of  the  nuclei  tuberis  laterales  in  Series  D  is 
practically  the  same  as  in  the  preceding  Series  AC,  Figs.  1  to  8.    Of 


Nuclei  Tuberis  Laterales  and  the  Ganglion  Opticum  Basale.    21 

the  two  fissures  referred  to  previously,  the  lateral  (just  medial  to  the 
pes  pedunculi)  does  not  occur,  since  the  obliquity  of  the  plane  of  section 
is  such  that  it  appears  only  in  sections  situated  further  caudally.  The 
medial  one  of  these  two  fissures,  however,  appears  in  every  section  from 
Figs.  11  to  15,  and  as  in  the  other  series,  forms  superficially  the  medial 
boundary  of  by  far  the  greater  portion  of  the  nuclei  tuberis  laterales. 

MACACUS   RHESUS. 

In  macacus  the  location  of  the  nuclei  tuberis  laterales  is  similar  to 
that  in  man,  except  that  in  most  sections  only  one  cell  group  is  shown, 
and  the  cell  mass  does  not  invade  the  medio-ventral  portion  of  the 
tuber.  A  comparison  of  Figs.  19  to  22  (macacus)  with  Figs.  1  to  8 
(man)  will  show  that  in  both  animals  the  nuclei  differ  but  little  as  to 
location.  The  two  parallel  furrows  on  the  base  of  the  brain  are  shown 
in  Figs.  19  and  20. 

In  both  man  and  macacus  the  nuclei  tuberis  laterales  have,  accord- 
ingly, practically  the  same  location,  and  by  far  the  greater  portion  is 
situated  in  the  previously  described  region;  this  region,  bounded  ven- 
trally  by  the  two  parallel  furrows,  laterally  by  the  pes  pedunculi,  and 
dorsally  by  the  fornix  column,  is  occupied  caudally  by  the  nucleus 
tubero-mammillaris,  and  the  nuclei  tuberis  laterales  displace  this 
nucleus  as  one  passes  orally,  and  after  the  optic  tract  has  fused  with 
the  tuber,  the  nuclei  tuberis  are  continued  orally  in  the  angle  between 
the  pes  pedunculi  and  the  optic  tract.  Since  the  purpose  of  this  paper 
is  in  part  to  give  a  foundation  for  experimental  study,  it  should  be 
noted  that  the  main  cell  mas^  of  fue  nuclei  tuberis  laterales  may  be 
accurately  loaded  in  the  intact  brain  just  beneath  the  surface  of  the 
most  lateral  portion  of  the  tuber  cinereum,  between  the  two  parallel 
furrows  and  just  caudal  (medial)  to  the  optic  tract.  In  toluidin  blue 
sections  the  nuclei  may  be  distinguished  as  circumscribed,  lightly 
staining  areas  by  means  of  the  unaided  eye.  As  previously  stated  no 
trace  of  the  nuclei  tuberis  laterales  is  present  in  the  cat,  while  in  the 
lemur  they  are  indicated  so  faintly  that  it  seems  best  not  to  attempt  to 
represent  them  in  the  illustrations. 

Separation  of  the  nuclei  tuberis  laterales  from  surrounding  cell  groups 
through  differences  in  cell  character. 
The  nuclei  tuberis  laterales  are  readily  distinguished  from  the  sur- 
rounding cell  groups  not  only  in  that  they  are  sharply  circumscribed 
cell  masses  having  a  constant  location,  but  also  through  their  charac- 


22  Edward  F.  Malone. 

teristic  cell  type.  A  reference  to  Fig.  36  will  serve  as  an  introduction 
to  the  further  consideration  of  the  differences  in  cell  character.  On 
comparing  the  cell  character  of  the  nuclei  tuheris  laterales  of  man  and 
macacus  (Figs.  39  and  45)  with  that  of  the  other  cell  groups  shown 
in  the  illustrations  it  will  at  once  be  evident  that  this  cell  type  differs 
radically  from  all  others  except  that  of  the  substantia  grisea  of  the 
third  ventricle  (Figs.  40  and  46)  ;  this  similarity  is  of  importance, 
and  its  significance  w'ill  be  considered  when  the  relationship  of  the 
various  cell  types  is  discussed.  Comparing  the  cell  type  of  the  nuclei 
tuberis  laterales  in  man  (Fig.  39)  and  in  maoacus  (Fig.  45)  it  is 
evident  that  aside  from  the  2>resence  of  pigmentation  in  man  (of  course 
not  to  be  expected  in  macacus)  there  is  no  essential  diiference.  The 
fundamental  characteristic  of  this  cell  type  in  both  forms  lies  in  the 
appearance  of  the  cytoplasm,  which  contains  an  extremely  small  amount 
of  Xissl  substance  in  the  form  of  fine  granules ;  the  relation  of  cell 
nucleus  to  cytoplasm  in  respect  to  volume  is  'also  practically  the  same, 
although  in  macacus  the  cytoplasm  is  relatively  less,  a  change  that  is 
to  be  expected,  as  will  appear  later.  In  describing  the  cells  of  the 
nucleus  ansae  peduncularis  (p.  18)  I  pointed  out  the  fact  that  if  two 
cell  groups  differ  as  to  the  amount  of  yellow  pigment  in  their  cells  they 
will  also  differ  in  other  respects,  and  that  homologous  groups  in  lower 
animals  (where  all  pigmentation  is  absent)  will  also  appear  different. 
Accordingly  the  presence  of  densely  packed  yellow  granules  in  the  cells 
of  the  nuclei  tuberis  laterales  of  man  is  not  the  only  distinctive  charac- 
teristic of  these  cells,  but  is  connected  with  other  characteristics, 
common  also  to  the  homologous  cells  of  macacus ;  these  characteristics 
involve  the  appearance  of  the  cytoplasm  and  make  it  possible  to  separate 
these  cells  without  difficulty  from  those  of  surrounding  groups.  If  we 
study  the  cell  type  of  the  substantia  grisea  in  all  four  fonns  (Figs.  40, 
46,  51,  and  56)  we  find  the  cell  type  practically  unchanged,  since  in  all 
fonns  the  cells  of  this  group  are  characterized  by  the  extremely  small 
amount  of  cytoplasm  and  relatively  large  nucleus;  in  many  cells  the 
cytoplasm  is  almost  absent,  and  one  sees  only  a  large  nucleus  and 
perhaps  one  or  more  cell  processes. 

Accordingly  the  cells  of  the  substantia  grisea  may  be  readily  separated 
from  those  of  all  portions  of  the  nuclei  tuberis  Laterales  not  only 
through  their  smaller  size,  but  also  through  the  minute  amount  of 
cytoplasm  in  relation  to  the  size  of  the  cell  nucleus;  however,  on  the 
border  line  between  these  two  types  of  cells  occur  occasionally  transi- 


Nuclei  Tuberis  Laterales  and  the  Ganglion  Opticum  Basale.     33 

tioD  types,  whose  significance  will  be  discussed  later.  The  nuclei 
tuberis  laterales  must  be  sliarply  distinguished  from  certain  regions  of 
the  substantia  grisea  in  whicli  the  cells  are  more  densely  packed 
together  than  in  other  portions  of  the  substantia  grisea.  Differences 
in  the  density  of  different  portions  of  the  substantia  grisea  may  be  seen 
in  many  sections,  and  very  definite  areas  of  closely  packed  cells  are 
shown  in  Figs.  17,  23  and  34.  While  in  such  dense  masses  the  cells 
are  possibly  somewhat  larger  than  the  remaining  cells  of  the  substantia 
grisea,  I  do  not  consider  the  difference  in  cell  type  clear  enough  to 
justify  one  in  setting  aside  such  dense  cell  masses  as  distinct  nuclei. 
Whatever  the  significance  of  these  denser  cell  masses  of  the  substantia 
grisea,  one  fact  is  certain :  they  diifer  both  in  location  and  in  cell 
type  from  the  nuclei  tuberis  laterales;  the  radical  difference  between 
these  two  kinds  of  cell  groups  should  be  clearly  recognized.  Of  course 
the  structure  of  the  cells  of  the  nuclei  tuberis  laterales  as  well  as  that 
of  the  cells  of  the  substantia  grisea  is  radically  different  from  that  of 
motor  cells,  while  the  small  size  of  both  types  of  cells  would  indicate 
that  the  connections  of  each  cell  were  very  limited;  this  correlation 
between  cell  size  and  the  extent  of  the  connections  of  each  cell  has 
already  been  referred  to. 

Eelationship  Of  The  Various  Cell  Groups  To  Oxe  Another. 

From  the  previous  description  of  the  various  cell  groups  together 
with  a  reference  to  the  figures  showing  their  cell  types,  it  will  be  evident 
that  the  cell  groups  herein  considered  fall  naturally  into  two  classes: 

1.  Those  composed  of  small  cells,  including  the  substantia  grisea, 
and  the  nuclei  tuberis  laterales,  and 

3.  Those  composed  of  large  cells,  including  the  basal  optic  gang- 
lion, the  nucleus  paraventricularis  hypothalami,  the  nucleus  tubero- 
mammillaris,  and  perhaps  also  the  nucleus  ansae  peduncularis  (gang- 
lion basale). 

Taking  up  the  first  class  of  nuclei  a  most  interesting  relation  will 
be  found  to  exist  between  the  substantia  grisea  and  the  nuclei  tuberis 
laterales.  The  cells  of  the  substantia  grisea  are  the  most  primitive 
of  the  hypothalamus.  The  constancy  of  cell  type  in  all  four  animal 
forms  would  suggest  this,  but  the  strongest  evidence  is  derived  from  the 
nature  of  the  cell  tyjie.  The  cell  type  of  the  substantia  grisea,  ^Wth  its 
scanty  cytoplasm,  resembles  that  of  embryonic  cells;  it  occurs  often  in 


24  Edward  F.  Malone. 

invertebrates ;  it  is  the  only  tj'pe  in  the  hypothalamus  which  approaches 
that  of  the  neuroglia  cells,  and  transition  forms  occur  concerning  which 
one  is  in  doubt  as  to  whether  they  are  neuroglia  cells  or  cells  of  the  sub- 
stantia grisea  ;  the  cells  are  most  abundant  near  the  ventricle  from 
whose  border  all  cells  of  the  hypothalamus  have  arisen,  while  laterally 
they  are  replaced  by  more  highly  differentiated  types  of  cells.  It  is  by 
no  means  certain  that  all  or  even  any  of  the  cells  of  the  substantia 
grisea  are  functional.  That  the  cells  of  the  nuclei  tuberis  laterales  have 
arisen  from  those  of  the  substantia  grisea  is  certain ;  if  we  look  in  the 
lemur  at  the  regions  where  in  higher  forms  the  nuclei  tuberis  laterales 
occur,  we  see  that  the  type  of  cell  is  somewhat  different  from  the  sur- 
rounding cells  of  the  substantia  grisea ;  this  difference  increases  in 
maeaeus,  and  still  more  in  man,  while  in  the  cat  there  is  no  indication  of 
any  new  group.  It  is  thus  evident  that  the  most  recent  cell  groups  of 
the  hypothalamus  (nuclei  tuberis  laterales)  have  arisen  directly  from 
the  oldest  cell  group  (substantia  grisea),  and  that  new  cell  groups  are 
not  necessarily  formed  by  the  further  histological  differentiation  of 
portions  of  cell  groups  which  are  already  highly  developed,  thus  result- 
ing in  a  very  highly  differentiated  cell  type,  but  that  on  the  contrary 
new  cell  groups  may  arise  from  cell  masses  which  have  remained  primi- 
tive. In  other  words  the  fact  tliat  a  cell  group  is  of  recent  origin  does 
not  give  any  information  as  to  the  extent  of  its  histological  differentia- 
tion, since  this  differentiation  depends  also  upon  the  development  of  the 
cell  group  from  which  the  new  group  arises. 

In  considering  the  second  class  of  cell  groups  in  the  hypothalamus, 
all  of  which  are  composed  of  large  cells,  the  nucleus  ansae  peduncularis 
requires  only  a  brief  consideration.  The  cell  t}-pe  of  this  cell  group 
differs  considerably  from  that  of  other  groups,  and  the  nucleus  is  prob- 
ably of  much  more  recent  phylogenetic  origin,  since  I  have  not  as  yet 
been  able  to  find  it  in  the  cat ;  in  order  to  consider  the  relations  of  this 
cell  group  a  study  of  the  neighboring  regions  of  the  telencephalon  will  be 
necessary,  and  I  shall  content  myself  with  having  shown  that  it  is  clearly 
distinguishable  from  the  basal  optic  ganglion,  and  with  having  given 
the  relative  location  of  both  groups  of  cells. 

Concerning  the  other  three  cell  groups  in  this  class  (basal  optic  gang- 
lion, nucleus  paraventricularis  hypothalami,  and  nucleus  tubero-mam- 
millaris),  the  phylogenetic  series  from  the  cat  to  man  is  far  too  short 
to  give  such  satisfactory  results  as  in  the  ease  of  the  nuclei  tuberis 
laterales,  since  the  former  three  groups  are  of  much  older  origin :  certain 


Nuclei  Tuberis  Laterales  and  the  Ganglion  Opticum  Basale.    25 

relationships  may,  however,  be  established.    Of  these  three  cell  groups 
the  basal  optic  ganglion  is  composed  of  cells  whose  type  is  by  far  the 
most  constant  in  the  different  animal  forms,  and  the  volume  of  this 
cell  group  is  reduced  in  the  lower  forms  to  a  less  extent  than  is  true  of 
the  two  other  nuclei;  moreover,  while  the  cell  tj-pe  of  the  basal  optic 
ganglion  approaches  closely  that  of  the  nucleus  paraventricularis.  there 
are  no  transition  forms  between  its  cell  type  and  that  of  the  nucleus 
tubero-mammillaris  or  that  of  the  substantia  grisea;  finally  the  basal 
optic  ganglion  is  almost  entirely  free  from  the  invasion  of  neighboring 
cells.    All  of  these  considerations  make  it  seem  probable  that  of  the 
three  nuclei  the  basal  optic  ganglion  is  the  oldest  and  most  highly 
developed.     Closely  related  to  the  basal  optic  ganglion  is  the  nucleus 
paraventricularis ;  it  is  more  reduced  in  size  in  the  lower  forms  than 
is  the  basal  optic  ganglion ;  between  its  cell  type  and  that  of  the  nucleus 
tubero-mammillaris  transition  forms  occur,  and  between  its  cells  lie 
many  cells  of  the  substantia  grisea.    While  these  facts  make  it  probable 
that  the  nucleus  paraventricularis  is  possibly  not  so  old  a  cell  group  as 
the  basal  optic  ganglion,  they  enable  one  to  state  positively  that  the 
nucleus  paraventricularis  is  related  on  the  one  hand  to  the  basal  optic 
ganglion  and  on  the  other  to  the  nucleus  tubero-mammillaris.  The  third 
cell  group,  the  nucleus  tubero-mammillaris.  is  like  the  nucleus  paraven- 
tricularis much  reduced  in  size  in  the  lower  animals ;  it  differs  strik- 
ingly from  both  the  other  cell  groups  in  being  not  sharply  circum- 
scribed, but  on  the  contran-  is  very  diffuse,"^  extending  through  a  large 
portion  of  the  h}-pothalamus ;  it  is  invaded  accordingly  by  the  cells  of 
the  substantia  grisea  to  a  much  greater  extent  than  is  the  nucleus  para- 
ventricularis.   Finally  the  cell  type  of  the  nucleus  tubero-mammillaris 
shows  a  transition  on  the  one  hand  to  that  of  the  nucleus  paraventric- 
ularis and  on  the  other  to  that  of  the  substantia  grisea.    Therefore  the 
nucleus  tubero-mammillaris,  while  related  to  the  nucleus  paraventricu- 
laris, is  related  to  the  substantia  grisea  in  three  ways :  transition  forms 
of  cells  occur,  the  cells  of  both  groups  are  intimately  intermingled,  and 
both  cell  groups  are  widely  and  diffusely  distributed.     The  nucleus 
tubero-mammillaris  is  therefore  the  least  highly  developed  of  the  three 
groups  (owing  to  its  relation  to  the  substantia  grisea),  but  whether  it  is 
also  the  most  recent  group  cannot  be  decided  here:  it  might  just  as 
well  be  an  old  and  relatively  stationary  group,  and  to  decide  this  point 
a  longer  phylogenetic  series  must  be  studied. 


26  Edivard  F.  M alone. 

The  relationship  of  these  various  cell  groups  of  the  hypothalamus 
may  be  summarized  as  follows : 

1.  The  substantia  grisea  ventriculi  tertii  is  the  oldest,  least  highly 
developed,  and  the  most  diffusely  distributed  cell  group  of  the  hypothal- 
amus. From  it  have  been  differentiated  at  least  two  (and  probably  all) 
of  the  four  cell  groups  that  lie  more  or  less  embedded  in  it.  Certain 
portions  of  the  substantia  grisea  appear  slightly  different  from  the 
rest,  in  that  the  cells  are  here  crowded  closely  together.  Whether  any  or 
all  of  the  cells  of  the  substantia  grisea  take  part  in  conducting  nervous 
impulses,  or  whether  this  group  serves  merely  as  material  from  which 
more  highly  differentiated  cell  groups  are  formed,  is  not  known. 

2.  From  the  substantia  grisea  at  least  two  different  cell  groups  have 
arisen ;  these  two  groujjs  of  cells  have  developed  along  diverging  lines. 
One  of  these  two  types  constitutes  the  nuclei  tuberis  laterales ;  it  is  by 
far  the  youngest  cell  group  in  the  hypothalamus,  the  first  indication  of 
its  presence  occurring  in  the  lemur.  The  second  cell  group  wiiich  has 
developed  from  the  substantia  grisea  is  the  nucleus  tubero^mammillaris ; 
its  cell  type  differs  radically  from  that  of  the  nuclei  tuberis  laterales, 
and  it  is  a  much  older  cell  group. 

3.  Along  the  same  line  as  the  nucleus  tubero-mammillaris,  and  to  a 
higher  degree  of  histological  differentiation,  have  developed  the  nucleus 
paraventricularis  and  the  basal  optic  ganglion  ;  the  nucleus  paraventric- 
ularis  is  intermediate  between  the  other  two  nuclei.  From  such  a 
short  phylogenetic  seines  as  that  from  man  to  the  cat  it  is  impossible  to 
determine  whether  all  three  of  these  nuclei  have  developed  iiidej)en- 
dently  from  the  substantia  grisea  or  whether  the  other  two  nuclei  liave 
developed  from  the  nucleus  tubero-mammillaris  and  thus  only  indirectly 
from  the  substantia  grisea. 

4.  A  more  extensive  series  of  animal  forms  should  enable  us  to  under- 
stand such  relations  better;  but  a  study  of  such  material  cannot  be 
expected  to  yield  satisfactory  results  unless  the  different  cell  groups 
be  distinguished  and  unless  the  question  of  the  relationship  as  ex- 
pressed by  the  cell  type  be  borne  in  mind. 

5.  Valuable  results  may  be  expected  also  from  a  study  of  the  histo- 
genesis of  these  various  cell  groups  as  revealed  in  the  later  stages  of 
development  of  the  human  brain;  little,  however,  is  to  be  expected 
from  such  material  unless  it  be  fixed  and  stained  witli  reference  to 
the  demonstration  of  cell  structure  in  the  nervous  system,  and  the 
methods  usually  employed  are  not  well  adapted  to  this  purpose. 


Nuclei  Tuberis  Laterales  and  tJie  Ganglion  Opticum  Basale.     27 

Conclusion. 

I  have  described  with  tlie  aid  of  illustrations  the  location  ^and  extent 
of  the  basal  optic  ganglion  and  nuclei  tuberis  laterales  in  four  forms 
of  manxiuals,  and  have  showii  their  relations  to  the  surrounding  cell 
groups  of  the  hypothalamus;  moreover  the  cell  types  of  these  two  cell 
groups  have  been  shown  to  differ  radically,  and  the  differences  in  cell 
character  have  been  pointed  out  through  which  one  can  readily  and 
accurately  distinguish  each  of  these  two  cell  groups  from  those  which 
surround  it.  Accordingly  the  principal  purpose  of  this  study  has  been 
accomplished,  namely,  to  prepare  a  foundation  of  such  a  character  as 
to  make  further  study  possible.  It  is  now  possible  to  study  in  the  same 
manner  neighboring  portions  of  the  telencephalon,  or  to  e.vtend  with 
relative  ease  the  present  study  of  the  hypothalamus  through  a  much 
longer  phylogenetic  series  than  the  critical  nature  of  this  study  has 
permitted ;  the  relations  of  the  various  cell  groups  of  the  hypothalamus 
can  be  studied  also  from  the  standpoint  of  histogenesis ;  or  the  nature 
of  the  cell  processes  and  the  character  of  the  nerve  endings  around  the 
cells  of  each  group  may  be  determined.  But  the  greatest  use  of  this 
study  is  that  it  makes  possible  intelligent  experimental  investigation 
of  the  hypothalamus.  Experimental  workers  have  been  inclined  to 
pass  over  differences  in  cell  character  and  to  make  little  or  no  attempt 
to  correlate  cell  function  with  cell  character,  and  fail  to  clearly  recognize 
a  fact  to  which  I  have,  in  a  previous  article,  called  attention  in  the 
following  words :  "  The  histological  character  of  a  nerve  cell  is  an  indi- 
cation of  its  function.  Differences  in  connections  with  portions  of  the 
organism  which  differ  merely  in  spatial  relations  do  not  involve  a 
difference  in  the  character  of  the  nerve  cells,  but  are  associated  merely 
with  the  location  of  the  nerve  cell ;  for  instance,  arm  and  leg  muscles, 
flexors  and  extensors  are  all  innervated  by  the  same  tyix;  of  cell, 
although  such  differences  in  peripheral  connections  correspond  to  the 
differences  in  the  position  of  the  corresponding  nerve  cells."  It  is  there- 
fore evident  that  experimental  work  which  detemiines  the  connections 
of  various  portions  of  a  given  region  with  different  portions  of  the 
organism,  without  taking  into  consideration  differences  of  cell  character, 
fails  to  distinguish  between  differences  of  connection  dependent  merely 
upon  spatial  differences  and  those  differences  of  connection  which 
involve  differences  in  cell  activity,  such  cell  activity  being  indicated, 
not  only  in  the  nervous  system  but  in  all  portions  of  the  entire  organism. 


28  Edward  F.  Malone. 

by  a  definite  type  of  cell  character.  That  these  two  aspects  of  the  func- 
tion of  the  nervous  system  might  be  successfully  studied  in  the  hypothal- 
amus by  the  use  of  experimental  methods  has  been  the  controlling 
factor  in  determining  the  nature  of  this  article. 

But  in  addition  to  preparing  a  foundation  for  exj3erimental  work  an 
attempt  has  been  made  to  partially  analyze  the  different  elements  in- 
volved in  the  histological  complex  of  each  cell  type  and  to  note  in 
homologous  cell  groups  of  various  animals  what  elements  of  cell  charac- 
ter are  constant ;  moreover  in  different  cell  groups  of  the  same  animal 
those  elements  of  the  entire  complex  of  cell  character  which  are  common 
to  the  cells  of  two  or  more  groups  have  been  noted,  and  by  this  means  an 
attempt  has  been  made  to  form  a  provisional,  although  crude,  idea  of  the 
degree  of  functional  relationship  among  the  different  cell  groups.  At 
first  sight  such  an  attempt  might  appear  premature,  or  it  might  even 
seem  as  though  the  task  of  the  histologist  were  finished  when  he  had 
pointed  out  the  existence  and  location  of  the  various  cell  groups,  and 
that  the  determination  of  the  functional  relations  of  different  cell  groups 
might  be  safely  left  to  the  experimental  worker.  Such  a  conclusion, 
however,  is  not  justified.  Many  regions  of  the  brain  are  so  complicated 
as  to  make  the  results  of  experimental  investigation  extremely  vague, 
and  in  more  suitable  regions  it  is  only  under  fortunate  conditions  that 
we  are  informed  of  the  functional  relations  of  a  cell  group  except  that  it 
plays  some  unknown  part  in  a  complex  mechanism  underlying  a  com- 
plex function.  In  solving  the  relations  of  the  different  components  of 
the  various  mechanisms  of  the  nervous  system  histology  must  play  an 
important  and  perhaps  the  principal  part,  since  in  the  nervous  system, 
as  in  the  rest  of  the  organism,  it  enables  us  to  locate  and  to  correctly 
state  the  function  of  every  cell  group  of  one  definite  cell  type,  if  the  func- 
tion of  this  cell  type  has  in  other  cell  groups  been  previously  clearly 
shown ;  that,  however,  it  is  not  self-sufficient  is  evident. 

But  the  neuro-histologist  has  a  much  more  difficult  task  than  that  of 
interpreting  in  a  general  way  the  differences  and  similarities  of  cell 
character  in  different  cell  types.  The  cell  types  of  certain  cell  groups 
resemble  one  another  through  possessing  certain  features  of  cell  char- 
acter common  to  all,  and  on  the  other  hand  differ  in  respect  to  other 
features  of  cell  character;  that  such  similarities  and  differences  of  cell 
type  are  an  indication  of  corresponding  similarities  and  differences  of 
function  I  have  already  clearly  shown  in  two  cases.  I  have  shown  (Am. 
Jour.  Anat.,  1913)  that  although  the  three  different  types  of  muscle  are 


Nuclei  Tuheris  Laterales  and  the  Ganglion  Opticum  Basale.     29 

supplied  by  three  distinct  types  of  nerve  cells  (p.  2),  these  three  types 
of  nerve  cells  possess  a  fundamental  similarity  of  cell  character ;  accord- 
ingly the  cell  type  varies  as  the  cell  function.  In  the  second  place  I 
have  shown  (Anat.  Rec,  1913)  that  all  cells  supplying  striated  muscle, 
whether  directly  (anterior  horn  cells,  etc.)  or  more  or  less  indirectly 
(large  pyramidal  cells,  cells  of  Deiters'  nucleus,  etc.)  possess  certain 
fundamental  features  of  cell  character  in  common.  This  fundamental 
character  makes  its  appearance  in  the  first  cell  of  any  efferent  chain 
which  is  set  aside  for  the  innervation  of  striated  muscle  and  is  present 
in  all  cells  of  such  a  series,  however  many  neurones  may  intervene 
between  the  cell  in  question  and  the  striated  muscle  fiber;  on  the  other 
hand  the  cells  in  such  a  voluntary  motor  chain  differ  in  relation  to  their 
position  in  the  chain.  It  is  thus  evident  that  the  particular  group  of 
elements  of  cell  type  which  corresponds  to  the  motor  function  of  the 
cell  may  be  distinguished  from  other  elements  of  cell  type  which  must 
correspond  to  the  other  known  or  unknown  influences.  The  feature 
of  cell  character  which  corresponds  to  motor  function  is  the  arrangement 
of  Nissl  substance  in  relatively  coarse,  discrete  bodies;  while  the  large 
size  (which  is  usually  supposed  to  be  common  to  motor  cells)  is  not 
directly  related  to  the  motor  function,  but  is  an  element  of  cell  character 
which  is  probably  (as  stated  previously)  an  indication  of  the  extensive 
connections  of  the  cell,  in  that  such  a  large  cell  receives  impulses  from 
(sphere  of  reception)  or  sends  impulses  to  (sphere  of  influence)  a  large 
territory. 

Accordingly  each  type  of  nerve  cell  should  be  regarded  not  as  distinct 
from  all  other  types,  but  as  the  result  of  different  influences  (due  to 
the  various  relations  of  the  nerve  cell  to  other  neurones  and  to  the  other 
portions  of  the  organism)  ;  when  so  viewed  the  manifold  variations  of 
cell  type,  including  cells  of  the  most  diverse  types,  as  well  as  those  of 
extremely  similar  types,  do  not  appear  to  offer  insuperable  difficulties, 
but  on  the  contrary  these  confusing  relations  of  cell  types  will  enable 
us  eventually  to  work  out  the  details  of  various  nervous  mechanisms. 
After  the  simultaneous  analysis,  by  the  aid  of  many  methods,  of  the 
histological  characteristics  and  the  functional  relations  of  difi'erent 
types  of  cells,  and  after  the  recognition  of  the  result  of  any  given 
functional  relation  upon  the  histological  character  of  the  cell,  the  neuro- 
histologist  will  be  able  to  take  a  further  step.  He  will  not  only  deter- 
mine one  portion  of  the  functional  relation  through  the  presence  of 
certain  features  of  cell  character  (which  is  already  possible  in  the  ease 


30  Edward  F.  Malone. 

of  various  types  of  motor  cells),  but  he  tvill  actttaUy  reconstruct  the 
greater  portion  of  the  entire  complex  of  cell  function  upon  the  basis  of 
the  presence  in  the  cell  of  certain  groups  of  histological  characters  the 
meaning  of  each  of  which  has  been  determined  independently  in  differ- 
ent cells.  A  beginning  has  already  been  made  in  that  we  are  able  to  dis- 
sociate in  motor  cells  the  characteristic  structure  (corresponding  to  the 
motor  activity)  and  the  cell  size  (corresponding  to  the  spheres  of 
influence  and  reception  of  the  cell)  ;  these  two  dissociated  elements  of 
cell  character  may  be  employed  independently  in  examining  any  type  of 
cell  in  the  entire  nervous  system.  Such  results  can  be  accomplished 
only  through  work  which  demands  the  exercise  of  the  greatest  critical 
ability,  but  that  they  will  be  attained  I  am  firmly  convinced. 

It  is  evident  that  no  such  results  are  yet  possible  in  the  case  of  the 
hypothalamus,  since  no  infonnation  is  available  concerning  the  func- 
tions of  any  of  its  cell  groups;  I  have  been  comixdled  therefore  to 
draw  very  general  conclusions  as  to  the  nearness  of  functional  relation- 
ship between  different  cell  groups,  since  these  conclusions  could  at 
present  be  based  merely  upon  greater  or  lesser  resemblance  of  cell  type. 
One  conclusion  as  to  the  function  of  these  cell  groups  is,  however, 
possible.  The  basal  optic  ganglion,  the  nuclei  tuberis  laterales,  the 
nucleus  paraventricularis  hypothalami  and  the  nucleus  tubero-mammil- 
laris  are  composed  of  cells  whose  histological  character  indicates  that 
they  are  not  efferent,  but  are  concerned  in  receiving  and  correlating 
incoming  impidses;  these  cells  do  not  possess  the  relatively  large,  dis- 
crete Nissl  bodies  characteristic  of  efferent  cells.  Moreover  (as  sug- 
gested by  Dr.  Donaldson)  the  small  size  of  such  cells  as  those  of  the 
nuclei  tuberis  laterales  would  seem  to  indicate  that  the  connections  of 
these  cells  were  less  extensive  than  those  of  the  cells  of  the  other  three 
nuclei.  But  now  that  these  various  cell  groups  have  been  described 
experimental  workers  will  be  in  a  position  to  attack  the  problems 
involving  the  connections  and  general  functional  significance  of  each 
of  these  cell  groups,  realizing  that  differences  of  cell  t}T)e  in  different 
cell  groups  cannot  be  ignored,  since  such  differences,  as  in  every-  portion 
of  the  organism,  correspond  to  differences  in  cell  function  ;  we  may  then 
expect  results  in  this  region  upon  which  may  be  based  in  turn  the  con- 
structive neuro-histology  outlined  above. 

To  those  who  prefer  to  employ  several  different  histological  methods 
I  shall  point  out  the  fact  that  the  Nissl  method  as  employed  by  me 
brings  out  very  clearly  the  differences  and  similarities  of  cell  type,  and 


Nuclei  Tuberis  Laterales  and  the  Ganglion  Opticum  Basale.     31 

accordingly  should  with  the  aid  of  experimental  work  and  of  other 
histological  methods  suifice  to  give  much  valuable  knowledge  as  to  the 
real  relations  involved  in  any  nervous  mechanism.  Every  method  of 
studying  the  nervous  system  has  its  advantages,  and  I  have  employed 
the  Nissl  method  merely  because  of  all  methods  it  is  the  most  valuable 
in  giving  a  complete  picture  of  the  cell  groups  of  the  entire  nervous 
system.  To  this  peculiar  advantage  of  this  method,  through  which  we 
can  compare  the  histological  character  of  all  cell  groups  of  the  nervous 
system,  I  should  like  to  direct  especial  attention.  Such  a  complete 
picture  of  the  cell  groups  of  the  nervous  system  would  seem  to  offer 
a  most  favorable  ground-work  for  assimilating  to  itself  isolated  facts 
observed  by  means  of  all  other  methods  of  studying  the  nervous  system ; 
without  other  methods  (histological,  experimental,  etc.)  it  is  of  course 
of  little  value.     (See  last  paragraph  of  introduction,  p.  3.) 

In  my  opinion  the  demands  of  such  work  are  so  exacting  that  each  in- 
vestigator who  attempts  to  gain  a  picture  of  the  cell  groups  of  a  large 
portion  of  the  nervous  system  should  confine  himself,  at  least  for  the 
most  part,  to  one  method,  and  of  all  methods  that  of  Nissl  shows  the 
cell  structure  in  the  clearest  manner.  It  is  not  isolated  cytological 
details  obtained  by  many  methods,  nor  is  it  the  proof  or  disproof  of  the 
actual  existence  in  the  living  cell  of  certain  features  of  cell  structure 
that  will  be  of  most  importance  in  the  above  outlined  constructive 
neuro-histology,  but  a  thorough  familiarity  with  the  location,  extent 
and  cell  type  of  the  different  cell  groups  of  practically  the  entire  nervous 
system,  together  with  a  most  critical  appreciation  of  the  differences, 
similarities  and  transitions  of  cell  type  of  all  these  cell  groups ;  without 
such  broad  knowledge  of  the  various  cell  groups,  which  must  of  course 
assimilate  to  itself  isolated  facts  regarding  the  function  and  the  connec- 
tions of  various  cell  groups  directly  and  indirectly  with  one  another,  and 
facts  regarding  tlie  number,  character,  and  mode  of  termination  of  the 
cell  processes  of  various  cell  types,  without  such  a  comprehensive  knowl- 
edge upon  which  to  build  we  could  not  ho]w  to  penetrate  far  into  the 
exact  relations  of  the  individual  neurones  of  the  nervous  system. 

LlTER.\TUl!E. 

The  literature  on  the  cell  groups  of  the  hypothalamus  has  been  of 
practically  no  assistance  to  me.     In  the  first  place  I  did  not  read  it 
until  I  had  worked  out  these  groups  in  the  human  brain,  and  upon 
3 


32  Edward  F.  Malone. 

reading  it  found  no  reason  for  modifying  my  conception  of  this  region. 
In  tile  second  place  some  of  tlie  articles  are  concerned  with  animal  forms 
so  far  removed  from  those  upon  which  I  have  worked  that  the  recogni- 
tion of  homologous  groups  is  by  no  means  certain.  Finally  the 
descriptions  of  cell  groups  are  often  so  inadequate  and  so  little  attention 
is  given  to  the  differences  in  cell  type  as  to  make  the  literature,  except  in 
rare  cases,  almost  unintelligible ;  this  is  true  especially  in  those  articles 
which  are  concerned  with  the  brain  of  man  and  the  higher  mammals. 
Accordingly  the  results  of  other  workers  on  the  cell  groups  of  the  hypo- 
thalamus have  not  been  incorporated  to  any  great  extent  into  the  body  of 
this  article,  but  are  here  appended  for  the  sake  of  completeness ;  to  treat 
the  greater  portion  of  the  literature  otherwise  would  render  the  subject 
of  this  article  needlessly  obscure.  The  following  consideration  of  the 
literature  is  given  for  the  benefit  of  those  who  wish  to  make  a  careful 
study  of  the  region  in  question ;  accordingly  it  appears  advisable  not  to 
give  an  abstract  of  the  various  articles,  since  the  complete  accounts  must 
be  read  by  those  for  whom  the  following  consideration  is  intended.  On 
the  contraiT  I  shall  for  the  most  part  confine  myself  to  a  consideration 
of  certain  points,  showing  wherein  certain  accounts  are  correct  and 
wherein  erroneous,  and  wherein  some  descriptions  are  altogether  unin- 
telligible. 

The  description  of  tlie  cell  groups  of  the  hypothalamus  contained  in  my 
monograph  on  the  human  diencephalon  has  been  referred  to  in  numerous 
places  in  the  text   and  a  further  reference  is   unnecessary. 

In  Strieker's  Manual  of  Histology  (American  translation)  Meynert 
gives  a  brief  but  fairly  clear  description  of  the  basal  optic  ganglion;  on 
p.  688  he  says:  "At  the  lateral  border  of  the  tuber  cinereum  lies  the 
inferior  optic  ganglion,  which  is  1.5  mm.  broad,  and  contains  spindle-shaped 
cells  30  /x  in  length  and  15  ii  in  breadth.  It  begins  just  above  the  optic 
commissure  and  stretches  along  immediately  over  the  tractus  opticus  as  far 
as  the  posterior  border  of  the  tuber  cinereum,  a  distance  of  more  than  a 
centimeter.  I  regard,  with  Luys,  this  optic  ganglion  as  a  part  of  the  tuber 
cinereum,  because  it  projects  downwards,  in  company  with  the  latter,  into 
the  lamina  cinerea,  beyond  the  surface  of  the  lamina  perforata  anterior,  of 
which  J.  Wagner  considers  it  to  be  a  part,  and  because  it  extends  farther 
backward  than  the  latter.  Like  the  tractus  itself,  however,  it  certainly 
follows  the  inner  border  of  the  anterior  pert,  space.  On  profile  sections 
(Fig.  270,  IT)  this  ganglion  has  a  sickle-like  shape,  the  concavity  looking 
forward.  According  to  Luys,  the  two  ganglia  touch  at  the  median  line, 
a  fact  which  I  have  not  been  able  to  verify,  etc."  From  this  description 
and  from  a  study  of  Fig.  268  it  is  clear  that  Meynert  had  in  mind  a  cell 
group  which  has  much  in  common  with  that  described  under  the  same  name 


Nuclei  Tuheris  Laterales  and  the  Ganrjlwn  Opticum  Basale.     33 

in  the  present  article.  The  exact  extent  is,  however,  not  clear,  and  we  have 
no  assurance  that  he  distinguished  between  this  cell  group  and  the  nucleus 
tubero-manimillaris,  or  that  he  included  all  portions  of  the  basal  optic 
ganglion.  Moreover  he  was  unable  to  observe  the  union  of  the  ganglia  of 
opposite  sides,  a  fact  which  makes  it  almost  certain  that  he  failed  to  include 
the  medial  portion  of  each  ganglion. 

When  we  attempt  to  understand  the  article  of  von  Lenhossek,  in  which 
he  divides,  in  man,  the  basal  optic  ganglion  of  Meynert  into  a  nucleus  supra- 
opticus  and  two  nuclei  tuheris,  we  are  confronted  with  an  impossible 
task.  We  are  informed  that  all  three  nuclei  are  composed  of  small,  spindle- 
shaped,  multipolar  nerve  cells,  as  well  as  neuroglia;  this  is  a  most  disturb- 
ing statement  and  renders  it  absolutely  impossible  to  form  any  idea  as  to 
what  cell  groups  the  author  had  in  mind.  For  by  no  means  can  the  cells 
of  the  basal  optic  ganglion  be  termed  small,  since  they  measure  up  to  35  m 
in  diameter,  and  compared  with  the  cells  of  the  nuclei  tuheris  and  the 
substantia  grisea  their  large  size  is  most  evident  (Fig.  37).  Accordingly 
the  word  "  small  "  would  seem  to  make  it  certain  that  Lenhossek's  nucleus 
supraopticus  cannot  be  a  portion  of  Meynert's  basal  optic  ganglion:  and 
yet  after  Meynert's  description  and  illustrations  it  would  seem  remarkable 
for  anyone  to  overlook  such  a  prominent  cell  group  as  the  oral  portion  of 
the  basal  optic  ganglion.  Again  we  are  informed  that  the  nucleus  anterior 
of  the  tuber  cinereum  is  the  largest  of  all  three  nuclei  (nucleus  supra- 
opticus and  the  anterior  and  postero-lateral  nuclei  of  the  tuber)  "  und 
Mldet  eigentUch  den  Hauptbestandteil  des  Tuber."  and  that  apparently  it 
reaches  the  median  line.  What  can  this  mean?  Can  the  "  small  "  cells  of 
this  anterior  nucleus  be  the  large  cells  of  the  nucleus  tubero-mammillaris, 
or  is  this  nucleus  a  portion  of  the  substantia  grisea?  One  is  almost  tempted 
to  infer  from  Lenhossek's  description  that  his  three  portions  of  Meynert's 
basal  optic  ganglion  are  a  complex  of  what  I  have  described  as  the  basal 
optic  ganglion  and  portions  of  the  nucleus  tubero-mammillaris,  all  of  which 
are  composed  of  large  cells,  and  that  the  cells  of  the  substantia  grisea  he 
may  possibly  have  considered  neuroglia;  since  he  employed  the  Weigert 
stain  such  a  mistake  is  by  no  means  improbable.  But  just  what  the  three 
nuclei  of  Lenhossek  represent  is  absolutely  impossible  to  determine. 

Kolliker's  description  of  the  nuclei  tuberis  and  the  basal  optic  ganglion  I 
have  in  a  previous  article  termed  vortrefflich ;  this  implies  a  uniform 
excellence  which  this  description  by  no  means  possesses.  It  were  more 
fitting  to  term  Kolliker's  account  of  these  nuclei  "  remarkable,"  for  it  is 
remarkable  as  to  insight  into  this  complicated  region,  and  at  the  same  time 
remarkable  for  the  contradictions,  incorrectly  labeled  figures,  and  other 
faults  which  detract  from  its  value.  Only  to  one  who  has  made  a  thorough 
study  of  this  region  is  the  difference  apparent  between  the  unintelligible 
description  of  von  Lenhossek  and  the  essential  excellence  of  that  of 
KoUiker,  and  from  1S96  until  1910  no  one  has  made  himself  sufficiently 
familiar  with  this  region  to  realize  the  many  excellent  points  in  Kolliker's 
work.     The  real  contributions  of  Kolliker  are  as  follows: 


34  Edward  F.  Malone. 

Es  miissen  'Nuclei  tuberis  von  den  Nuclei  supraoptici  unterschieden  wer- 
den  (p.  602) . .  .Die  Nuclei  tuberis  kommen  mehr  in  den  medialen  Gegenden 
vor,  besitzen  kleine  Nervenzellen,  etc.  (p.  603) . .  .Die  Nuclei  supraoptici  ha- 
ben  grossere  Zellen,  etc.  (p.  603).  Moreover  Figs.  704  and  705  are  correctly 
labeled  and  entirely  intelligible.  Fig.  702  shows  the  nuclei  tuberis  laterales 
excellently,  but  unfortunately  they  are  termed  "ganglia  optica  basalia"; 
on  p.  599  these  groups  are  correctly  termed  nuclei  tuberis,  while  on  p.  519 
they  are  incorrectly  termed  (in  text)  "ganglia  optica  basalia."  This 
uncorrected  error  in  Fig.  702  is  unfortunately  characteristic  of  the  incon- 
sistent nature  of  Kolliker's  description.  But  in  Fig.  631  we  meet  a  further 
difficulty, since  the  cell  mass  of  the  tuber  which  in  tig.  631  (rabbit)  is  termed 
"  Ganglion  opticum  basale  "  is  stated  on  p.  603  to  represent  the  "  Ganglia 
tuberis";  in  reality  this  cell  mass  is  simply  a  portion  of  the  substantia 
grisea  and  is  not  the  nuclei  tuberis  laterales  (which  are  shown  in  Figs. 
702,  G.  o.  b.,  and  705,  G.  t).  On  p.  599  he  has  incorrectly  stated  that  the 
nuclei  tuberis  are  continued  more  or  less  definitely  into  two  of  the  groups 
of  the  corpus  mammillare,  nucleus  intercalatus  (or  possibly  a  separated 
portion  of  the  medial  nucleus  as  discussed  in  my  monograph  on  the 
diencephalon)  and  the  nucleus  tubero-mammillaris;  this  proves  beyond 
doubt  that  Kolliker's  nuclei  tuberis  are  to  be  regarded  as  topographical 
groups  rather  than  determined  by  a  definite  cell  character.  On  the  other 
hand  he  has  confused  a  portion  of  the  nucleus  tubero-mammillaris  with 
the  basal  optic  ganglion;  on  p.  600  he  says:  Dieser  Kern  ist  schon  in  der 
Fig.  703  in  der  Grenzlage  zwi-schen  dem  Tract  us  opticus,  dem  Reste  des 
Pes  pedunculi  und  dem  Nucleus  Tuberis  lateralis  in  den  ersten  Anfdngen 
vorhanden  und  zwar  me  in  der  Linsenkernschlinge  drin,  wird  aber  nach 
vorn  immer  grosser,  etc.  Accordingly  Kolliker  has  included  portions  of 
the  nucleus  tubero-mammillaris  with  the  nuclei  tuberis  on  the  one 
hand,  and  on  the  other  with  the  basal  optic  ganglion.  The  vacant  space 
surrounding  the  third  ventricle  in  his  figures  raises  the  question  as  to  how 
he  regarded  the  substantia  grisea  of  the  third  ventricle;  its  relations  to  the 
other  nuclei  he  has  failed  to  point  out,  and  in  how  far  he  confused  this 
cell  mass  with  the  nuclei  tuberis  we  cannot  determine,  except  in  Fig.  631 
in  case  of  the  rabbit,  where  this  error  is  apparent;  this  cell  group  of  the 
rabbit  is  the  substantia  grisea  and  does  not  correspond  to  the  nuclei  tuberis 
as  shown  in  his  figures  of  man.  In  conclusion,  Kolliker's  description  and 
figures  are  often  very  faulty,  but  on  the  other  hand  in  his  figures  may  be 
recognized  in  many  cases  cell  groups  correctly  represented. 

Cajal  (pp.  756-757)  describes  in  rodents  a  cell  group  under  the  name 
ganglia  peri-kiasmatico  o  tangencial  which  without  doubt  is  homologous 
with  the  basal  optic  ganglion  of  higher  mammals.  He  is  in  doubt  as  to 
whether  it  corresponds  to  the  supraoptic  nucleus  of  Lenhossek  and  the 
supraoptic  complex  of  Kolliker.  As  previously  explained  it  is  impossible  to 
identify  the  supraoptic  nucleus  of  Lenhossek,  especially  since  he  says  that 
it  is  composed  of  "  small  "  cells;  the  description  of  Cajal  as  well  as  his 
excellent  illustration   (Fig.  640)  shows  that  the  ganglio  peri-kiasmiitico  is 


Nuclei  Tuheris  Laterales  and  the  Ganglion  Opticum  Basale.     35 

composed  of  large  cells.  Cajal  further  states  that  there  is  a  great  difference 
in  form,  position  and  extent  between  this  ganglion  of  rodents  and  the  above 
human  centers.  While  I  have  not  yet  studied  the  basal  optic  ganglion  in 
rodents  I  am  strongly  of  the  opinion  that  this  ganglion  in  rodents  does  not 
differ  essentially  either  in  form,  position  or  extent  from  the  basal  optic 
ganglion  in  man;  I  base  this  conclusion  on  the  constancy  of  this  ganglion 
from  man  to  the  cat,  and  upon  Cajal's  description  and  illustration.  There 
is  no  difficulty  in  recognizing  that  the  centers,  which  Cajal  describes  in  the 
tuber  under  the  names  of  anterior,  posterior  and  superior  nuclei  of  the 
tuber,  are  portions  of  the  substantia  grisea  and  have  nothing  to  do  with 
the  nuclei  tuberis  laterales  of  man  and  macacus;  the  difference  in  loca- 
tion of  these  three  nuclei  and  the  fact  that  there  is  every  reason  to  believe 
that  the  nuclei  laterales  are  not  present  in  the  rodents  (appearing  first 
in  the  lemur)  renders  confusion  impossible.  Under  the  name  niicleo 
subventricular  Cajal  describes  (p.  731)  a  cell  column  which  is  beyond 
doubt  homologous  with  the  nucleus  paraventricularis  hypothalami,  which 
I  have  described  in  man  and  the  higher  mammals;  Cajal's  description  and 
illustration  (Fig.  604,  T)  place  this  fact  beyond  the  possibility  of  a  doubt. 
Accordingly  Cajal's  description  of  the  hypothalamus  enables  us  to  recognize 
positively  two  characteristic  cell  groups  in  rodents  one  of  which  is  homolo- 
gous with  the  basal  optic  ganglion  and  the  other  with  the  nucleus  paraven- 
tricularis hypothalami  of  higher  animals.  Concerning  the  complex  of  the 
nucleus  tubero-mammillaris  we  are  left  in  doubt. 

The  presence  of  the  basal  optic  ganglion  and  the  nucleus  paraventricularis 
is  certain  not  only  in  rodents  but  even  in  the  marsupials,  as  is  seen  from 
Ziehen's  description  of  the  brain  of  Pseudochirus  peregrinus.  In  describing 
Fig.  28  he  says  (p.  713) :  Sehr  schim  ausgepriigt  ist  beiderseits  das  Gang- 
lion opUciim  basale  (nucleus  supraopticus).  Rechts  sendet  es  einen  lang- 
gestreckten  AusUiufer  in  das  Pedamentum  laterale.  Ingesammt  erstreckt 
es  sich  ilber  1.3  mm.  in  sagittaler  Richtung.  Seine  Zellen  messen  21  fi. 
Frontalwiirts  reicht  es  noch  ein  wenig  iiber  den  vorderen  Chias-marand 
hinaus.  Einen  Zerfall  in  mehrere  Zellgruppen,  wie  ihn  Lenhossck  und 
Kolliker  bei  dem  Menschen  beschrieben  haben.  vermijchte  ich  nicht  sicker 
nachzuweisen.  On  p.  714  in  describing  this  same  section  (Fig.  28)  Ziehen 
says:  In  der  grauen  Masse  zu  beiden  Seiten  des  3  Ventrikcls  kann  man — in 
der  Reihenfolge  von  untcn  nach  oben — folgende  The.ile  unterscheiden. 
Vnmittelbar  oberhalb  des  Chiasmas  folgt  der  kleinzellige  Nucleus  tuberis 
(evidently  the  substantia  grisea),  auf  diesen  ein  gross-zelUger  lateraler 
Kern,  in  icelchem  der  Fornixsiiule  eingebettet  ist  (a  portion  of  my  nucleus 
tubero-mammillaris),  und  ein  eben  so  grosszelUger  medialer  Kern,  welcher 
der  Yentrikelwand  unmittelbar  anliegt.  This  latter  nucleus  is  evidently 
homologous  with  the  nucleus  paraventricularis  hypothalami,  and  I 
thoroughly  agree  with  Ziehen  when  he  continues  as  follows:  Bei  der  sehr 
starken  Auspriigung  des  letzeren  war  ich  sehr  erstaunt,  eine  iihnliche 
Bildung  bei  anderen  Siiugern  nirgends  beschrieben  zu  linden.  Ich  mil  den 
Kern    einstweilen    als    Nucleus    subcommissuralis    bezeichnen.      Ziehen's 


36  Edward  F.  M alone. 

illustrations  unfortunately  show  almost  nothing  ot  the  various  cell  groups, 
but  his  concise  description  is  most  valuable. 

Worthy  of  special  notice  is  the  valuable  contribution  of  Friedemann 
entitled  Die  Cytoarchitektonik  des  Zivisvhenhirns  der  Cercopitheken  mit 
besonderer  Beriicksichtigung  des  Thalamus  opticus.  As  the  title  indicates 
more  attention  is  paid  to  the  thalamus  than  to  the  hypothalamus,  although 
both  the  pars  mammillaris  and  the  pars  optica  are  by  no  means  neglected. 
In  this  article  Friedemann  takes  into  consideration  my  monograph  on  the 
human  diencephalon.  To  consider  intelligently  Friedemann's  account  of  the 
cell  groups  of  the  hypothalamus  it  is  necessary  to  realize  that  his  chief 
purpose  is  to  subdivide  this  region  into  topographical  cell  groups,  which 
are  not  necessarily  composed  exclusively  ot  one  type  of  cell,  but  may  repre- 
sent complex  centers  formed  by  the  overlapping  of  several  primary  nuclei; 
on  the  other  hand  he  has  occasionally  divided  a  cell  mass,  composed  of  cells 
of  the  same  type,  into  subgroups  whenever  factors  other  than  cell  character 
(such  as  number  and  grouping  of  cells)  would  permit.  In  this  he  has  con- 
sistently endeavored  to  give  us  a  topographical  subdivision  into  as  many 
areas  as  may  possibly  be  distinguished  in  cell  preparations.  It  is  accord- 
ingly evident  that  my  cell  groups  (primary  nuclei i  have  been  set  aside  in 
virtue  of  the  possession  of  one  type  of  cell  which  corresponds  to  some  known 
or  unknown  cell  function  or  functions  and  that  the  topographical  value  of 
my  work  is  primarily  concerned  with  the  location  and  distribution  of 
such  a  single  cell  type;  and  on  the  other  hand  it  is  evident  that  Friede- 
mann's subdivision,  not  being  confined  to  this  one  criterion  of  cell  character, 
results  in  a  much  more  minute  subdivision  which  is  of  great  value  in  exact 
orientation,  and  that  his  cell  groups  may  represent  primary  nuclei,  portions 
of  such  nuclei,  or  complex  nuclei,  according  to  the  criteria  employed  in  the 
establishment  of  each  cell  group.  Apparently  Friedemann  has  not  grasped 
this  fundamental  difference  in  our  aims,  since  (p.  312)  he  says:  So  xvichtig 
auch  u.  a.  die  von  Malone  festgesteijte  Tatsaehc  ist,  dass  im  Thalamus 
opticus  keine  ZeUen  vom  viotorischen  Typus  vorkomm,en,  so  sehr  wir  auch 
in  vielen  Einzelheiten  mit  diesem  Autor  iibereinstimmen,  als  topographis- 
ches  Einteilungsprinzip  konnen  wir  seine  Methode  nicht  anerkennen. 
It  is  certainly  my  earnest  desire  that  no  one  should  consider  the  method 
I  have  employed  as  a  topographisches  Einteilungsprinzip.  since  I  myself 
have  never  so  considered  it,  and  had  supposed  that  I  had  made  this  fact 
evident.  But  Friedemann's  work  is  something  more  than  a  pure  topo- 
graphical subdivision,  since  he  has  informed  us  as  to  whether  a  cell  group 
has  been  recognized  on  account  of  its  cell  type,  or  whether  it  is  differen- 
tiated from  the  surrounding  cells  on  the  basis  of  other  criteria;  in  other 
words  he  has,  in  giving  us  this  minute  and  most  valuable  topographical 
subdivision,  not  lost  sight  of  the  distribution  of  the  various  cell  types.  The 
fact  that  Friedemann  and  myself  have  subdivided  the  diencephalon  from 
different  points  of  view  is  a  fortunate  circumstance,  and  the  difference  in 
results  is  to  be  expected  and  welcomed.  Other  subdivisions  from  other 
standpoints  are  desirable,  but  it  is  essential  that  we  be  accurately  informed 


Nuclei  Tuberis  Laterales  and  the  Ganglion  Opticum  Basale.     37 

just  what  criteria  have  been  employed  in  setting  aside  each  cell  group:  and 
moreover  we  should  carefully  distinguish  between  such  cell  groups  of 
different  authors  as  are  identical  and  such  as  do  not  entirely  correspond. 

Taking  up  the  various  cell  groups  of  the  hypothalamus,  Friedemann  des- 
cribes the  nucleus  intercalatus  corporis  mammillaris  and  the  nucleus 
paraventricularis  just  as  I  had  found  them  in  man,  and  he  adopts  my  names 
for  them.  He  also  adopts  my  name  "nucleus  mammillo-infundibularis  "; 
I  have  previously  in  this  article  pointed  out  that  this  name  must  be 
changed,  and  have  adopted  instead  that  of  nucleus  "  tubero-mammillaris." 
But  it  is  not  clear  to  me  that  Friedemann  has  included  all  the  cells  in  this 
group  which  I  have  assigned  to  it;  unfortunately  the  illustrations  (photo- 
graphs) are  of  little  value  in  showing  the  distribution  of  each  cell  type. 
It  is  certain  that  his  nucleus  posterior  pedamenti  lateralis  (Fig.  9,  Pip) 
is  a  portion  of  my  nucleus  tubero-mammillaris;  moreover  Friedemann 
has  called  attention  to  the  fact  (p.  367)  that  his  lamina  grisea  separans 
(Fig.  9,  Igs)  is  a  portion  of  this  same  nucleus.  Such  subdivisions  are  of 
course  desirable  for  their  topographical  value,  but  the  common  cell  charac- 
ter should  be  kept  in  mind.  In  considering  the  basal  optic  ganglion  Friede- 
mann has  fallen  into  the  same  error  that  I  did  in  my  monograph  on  the 
diencephalon,  and  for  the  same  reason,  since  both  of  us  were  not  primarily 
interested  in  the  telencephalon;  this  error  consists  in  considering  merely 
the  cell  mass  which  follows  the  oro-lateral  border  of  the  optic  tract  as 
constituting  this  ganglion.  It  is  certain  that  to  the  basal  optic  ganglion  of 
cercopithecus  must  be  added  Friedemann's  "  nucleus  anterior  pedamenti 
lateralis  "  (p.  370  and  Fig.  11,  Pla)  as  well  as  his  "  stratum  supraopticum  " 
(p.  369  and  Fig.  9,  Sso.).  That  these  two  cell  groups  belong  to  the  basal 
optic  ganglion  is  clear  both  from  the  descriptions  and  illustrations.  It  is 
unfortunate  that  neither  the  cross  sections  nor  the  horizontal  sections 
illustrated  give  the  region  just  caudal  to  the  optic  chiasm.  Friedemann's 
cross  sections  may  be  best  compared  with  my  Series  D  of  man  (Figs.  11  to 
18)  since  the  plane  of  section  is  similar:  in  comparing  these  two  series  it  is 
evident  that  the  most  oral  section  (Fig.  11)  of  Friedemann  stops  short 
of  a  most  important  region  of  the  basal  optic  ganglion.  Again  his  first 
horizontal  section  (Fig.  12)  likewise  fails  to  give  this  region  near  the  optic 
chiasm.  (The  omission  of  this  region  is  only  natural,  since  Friedemann's 
article  deals  primarily  with  the  diencephalon.)  Accordingly  I  see  no  reason 
to  believe  that  the  basal  optic  ganglion  in  cercopithecus  is  essentially 
different  from  that  in  other  animals  from  man  to  the  marsupials. 

In  Friedemann's  description  of  cercopithecus  I  am  unable  to  find  any 
cell  groups  which  would  correspond  to  the  nuclei  tuberis  laterales, 
and  unfortunately  the  illustrations  are  of  such  a  nature  (photographs)  as 
not  to  reveal  the  presence  of  such  groups.  It  is  not  probable  that  the  nuclei 
tuberis  laterales  are  wanting  in  cercopithecus,  since  they  are  present  in 
macacus  and  are  indicated  even  in  the  lemur;  their  presence,  if  indeed  they 
occur  in  cercopithecus,  might  easily  be  overlooked  unless  one  had  seen 
them  in  man,  where  these  nests  of  cells  are  very  definite.    The  fact,  which 


38  Edward  F.  Malone. 

Friedemann  himself  has  pointed  out  (p.  367-8),  should  be  most  carefully 
noted:  Friedemann's  nuclei  of  the  tuber  cinereum  are  different  portions 
of  the  substantia  grisea;  he  calls  attention  moreover  to  the  fact  (p.  367) 
that  the  substantia  grisea  has  been  divided  into  these  nuclei  rather  on 
account  of  topographical  relations  and  differences  in  the  crowding  together 
of  cells  than  on  account  of  differences  in  cell  form.  Accordingly  these 
topographical  cell  groups  of  the  substantia  grisea  must  not  be  confused 
with  the  nuclei  tuberis  laterales,  since  the  latter  are  composed  of  cells  of 
another  type. 


Bibliography. 

Cajal,  S.  R.  y:  Textura  del  sistema  nervioso,  etc.,  1904,  vol.  2. 
FoREL,    A. :      Beitrage    zur    Kenntnis    des    Thalamus    opticus,  etc. 
Gesammelte    hirnaiiatomische    Abhandlungen,    1872,    pp.    17-43. 

(Published  originally  in  Bd.  66  der  Sitzb.  der  k.  Akad.  d.  Wis- 
sensch.  in  Wien,  dritte  Abteilung,  Juni.) 
Friedemaxn,  M.  :    Die  Cjtoarchitcktonik  des  Zwischenhirns  der  Cer- 

copitheken,  etc.     Jl.  f.   Psychologie  u.   Neurol.,   1911,   XVIII, 

Erganzungsheft  2. 
Ganser,  L.  :    Yergleichende  anatomische  Studien  iiber  das  Gehirn  des 

Maulwurfs;  Morpholog.  Jahrbuch,  1882,  VII. 
KoLLiKEE,  A. :    Handbuch  der  Gewebelehre,  1896,  II. 
VON  Lenhossek,  M.  :  Beobaclitungeu  am  richirn  des  Menschen,  Anat. 

Anz.,  1887,  II,  Nr.  14. 
Malone,  E.  :    Uber  die  Kerne  des  menschlichen  Diencephalon.    .\us 

dem  Anhang  zu  den  Abhandlungen  der  konigl.  preuss.  Akad.  d. 

Wissensch.,  1910. 

Observations  concerning  the  comparative  anatomy  of  the  dien- 
cephalon.   Anat.  Eec,  1912,  VI,  No.  7. 

Eecognition  of  members  of  the  somatic  motor  chain  of  nerve 

cells,  etc.    Anat.  Kec,  1913,  VII,  No.  3. 

The  nucleus  cardiacus  nervi  vagi,  etc.    Am.  Jour.  Anat.,  1913, 

XV,  No.  1. 
JIeyxert,  Th.:     The  brain  of  mammals.     Strieker's  manual  of  his- 
tology (American  translation),  1872. 
Ziehen,  Th.  :    Das  Centralnervensystem  der  Monotremen  und  Mar- 

supialier.    Teil  2,  erster  Abschnitt.     From  Semon's  Forschungs- 

reisen.  III. 


Nuclei  Tuberis  Laierales  and  the  Ganglion  Opticum  Basale.     39 

Explanation  of  Illustrations. 

Figs.  1  to  35  and  lA  to  35A  have  been  prepared  in  the  following  manner: 
The  entire  section  was  in  each  case  projected  at  a  magnification  of  ten 
diameters,  and  the  outlines  together  with  as  many  details  as  possible  were 
drawn  in  pencil;  even  bloodvessels,  tears  in  the  section,  particles  of  detritus, 
etc.,  were  drawn  in  so  as  to  serve  the  purpose  of  orientation.  Under  the 
microscope  at  the  same  magnification  (ten  diameters)  the  drawing  was 
completed,  with  the  occasional  aid  of  higher  magnification.  (Of  course  the 
drawings  were  not  begun  until  I  had  a  clear  idea  of  the  location,  extent, 
cell  character,  and  relations  of  the  various  cell  groups.)  This  completed 
drawing  is  not  used  for  publication  but  serves  as  the  basis  for  two  others. 
In  the  first  place  an  outline  drawing  is  made  (by  tracing)  and  the  various 
structures  labeled  (Figs.  lA  to  35 A) ;  in  these  drawings  a  rectangular  area 
is  indicated  which  contains  the  region  in  which  we  are  interested.  A  second 
series  of  drawings  (Figs.  1  to  35)  are  traced  from  that  portion  of  the  original 
included  within  the  rectangular  area,  and  the  details  filled  in,  under  control 
of  the  microscope.  Finally  these  two  series  of  drawings  have  been  repro- 
duced in  parallel  columns;  in  order  to  do  this  the  outline  drawings  have 
been  more  greatly  reduced  in  size  than  the  detailed  drawings,  the  resulting 
magnification  being  indicated  in  the  explanation  of  each  plate.  Figs.  1  to  IS 
have  been  so  much  reduced  that  they  should  be  examined  with  the  aid  of  a 
hand  lens;  the  cells  of  the  nuclei  tuberis  laterales  are  especially  difficult  to 
recognize  without  such  aid. 

It  is  important  to  understand  just  what  these  drawings  are  intended  to 
illustrate.  The  outline  series  (Figs.  lA  to  35A)  needs  no  explanation,  except 
the  statement  that  the  size  and  relations  of  the  structures  shown  have 
been  accurately  reproduced.  In  the  other  series  (Figs.  1  to  35)  the  distri- 
bution of  each  cell  group  is  accurately  shown,  each  being  indicated  by  one 
type  of  cell  which  may  be  distinguished  in  all  drawings  from  the  cells  of 
all  other  cell  groups;  thus  different  cell  groups  may  be  distinguished  when 
their  cells  are  intermingled,  and  it  is  thereby  possible  to  reproduce  the 
actual  extent  of  each  cell  type  without  resorting  to  an  arbitrary  boundary. 
On  the  other  hand  in  order  to  indicate  differences  in  cell  type  it  has  been 
necessary  to  make  the  cells  much  larger  than  they  appear  at  such  a  low 
magnification,  and  this  in  turn  involves  a  reduction  in  the  number  of  cells. 
However,  the  relative  density  of  each  cell  mass  in  its  various  portions  is 
shown,  and  the  relative  size  of  different  cell  types  has  been  maintained, 
although  this  difference  in  size  has  in  some  cases  been  somewhat  exag- 
gerated in  order  to  make  it  more  obvious.  For  a  true  picture  of  each  cell 
type  one  must  study  Figs.  38  to  58,  in  which  each  cell  type  has  been 
reproduced  exactly  as  it  appears. 

The  exact  cell  size  and  distance  between  the  cells  of  the  nuclei  tuberis 
laterales  and  the  basal  optic  ganglion,  as  well  as  that  of  the  surrounding 
cell  groups,  has  been  reproduced  exactly  in  Figs.  36  and  37,  which  also  show 
the  different  cell  types  as  they  appear  at  a  relatively  low  magnification. 
These  two  illustrations  (Figs.  36  and  37)  will  be  explained  on  the  pages 
facing  them. 


40  Edward  F.  Malone. 

Figures  3S  to  58  were  drawn  as  follows: 

With  the  aid  of  the  camera  lucida  the  cell  outlines  and  as  many  details 
as  possible  were  drawn  in  pencil.  Then  under  control  of  the  microscope 
the  exact  appearance  of  the  cells  was  reproduced  in  water  color.  Each 
cell  is  to  be  regarded  as  an  individual  and  the  spatial  relations  of  the  cells 
in  each  figure  do  not  correspond  to  those  between  the  cells  of  the  corres- 
ponding cell  group  in  the  preparations.  In  these  figures  the  cells  have 
been  reproduced  by  combining  to  a  certain  extent  different  optical  sections 
of  the  same  cell.  All  cells  have  been  drawn  at  a  magnification  of  580  diame- 
ters, and  have  been  reproduced  without  reduction.  Figs.  38  to  58  were  all 
drawn  from  cross-sections  of  the  brains  of  the  various  animals.  In  Series 
D  of  man  the  plane  of  section  differs  widely  from  that  of  a  cross-section, 
and  yet  (without  making  a  careful  study  of  this  point)  I  was  unable  to 
observe  that  this  difference  in  the  plane  of  section  was  correlated  with  any 
difference  in  the  appearance  of  the  cell  types  here  illustrated.  On  the  con- 
trary I  can  state  positively  that  each  of  the  above  types  of  cells  was  as 
characteristic  in  Series  D  as  in  Series  AC,  and  that  differences  in  the  plans 
of  section  could  not  possibly  lead  to  a  confusion  of  the  different  cell  types. 


42  Edward  F.  Malone. 


PLATE  I. 

Homo.     Seeies  AC. 

Magnification  of  Figs.  1  to  4  =  5  diameters. 
Magnification  of  Figs.  lA  to  4A  =  2  diameters. 

•  =  Cells  of  the  nucleus  tubero-mammillaris  (formerly  known  as  nucleus 

mammillo-infundibularis) . 

•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 

A  =  Cells  of  the  nuclei  tuberis  laterales  (n.  t.  1.).     (Examine  with  baud 

lens ) . 
O  =  Cells  of  the  ganglion  opticum  basale  (g.  o.  b.). 
Q  =  Cells  of  the  corpus  hypothalamicum    (Luysii). 


THE  JOHNS   HOPKINS  HOSPITAL   REPORTS. 


PLATE   I. 


4A  Edward  F.  Malone. 


PLATE  II. 

Homo.     Series  AC    (Continued). 

Magnification  of  Figs.  5  to  7  =  5  diameters. 

Magnification  of  Figs.  5A  to  7A  =  2  diameters. 

•  =  Cells  of  the  nucleus  tubero-mammillaris  (nucleus  mammillo-infundib- 

ularis). 
•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 
A  =  Cells  of  the  nuclei  tuberis  laterales  (n.  t.  1.).     (Examine  with  hand 

lens). 
O  =  Cells  of  the  ganglion  opticum  basale  (g.  o.  b.). 
O  =  Cells  of  the  corpus  hypothalamicum    (Luysii). 


THE  JOHNS   HOPKINS  HOSPITAL   REPORTS. 


PLATE   II. 


Fio.  7. 


Fig.  7.\. 


46  Edward  F.  Malone. 


PLATE  III. 

Homo.     Series  AC   (Co.ncluded). 

Magnification  of  Figs.  8  to  10  =  5  diameters. 
Magnification  of  Figs.  8A  to  lOA  =  2  diameters. 

{Cells  of  tlie  nucleus  tubero-mammillaris  (nucleus  mammillo-infun- 
dibularis). 
Cells  of  the  nucleus  paraventricularis  hypothalami. 
•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 
A  =  Cells  of  the  nuclei  tuberis  laterales  (n.  t.  1.).     (Examine  with  hand 

lens ) . 
O  =  Cells  of  the  ganglion  opticum  basale  (g.  o.  b. ). 

O  =  Cells  of  the  nucleus  ansae  peduncularis  (ganglion  basale  of  Kolliker). 
Note  that  the  cells  of  the  nucleus  tubero-mammillaris  and  those  of  the 
nucleus  paraventricularis  are  represented  by  the  same  type  of  cell;  more- 
over the  same  type  of  cell  has  been  employed  to  represent  the  cells  of  the 
corpus  subthalamicum  and  those  of  the  nucleus  ansae  peduncularis.  It 
seems  advisable  to  use  one  type  of  cell  to  represent  the  cells  of  two  different 
groups,  when  these  two  groups  are  clearly  distinguishable  topographically, 
rather  than  to  multiply  the  number  of  symbols  used,  and  thus  render  it 
more  diflBcult  to  distinguish  between  them.  The  outline  drawings  will 
make  a  confusion  of  such  cell  groups  impossible. 


THE  JOHNS  HOPKINS  HOSPITAL  REPORTS. 


PLATE  III. 


Fig.  9. 


Fig.  9a. 


Fig.  10. 


Fig.  10a. 


48  Edward  F.  Malone. 


PLATE  IV. 

Homo.     Series  D. 

Magnification  of  Figs.  11  to  14  =  5  diameters. 
Magnification  of  Figs.  IIA  to  14A  =  2  diameters. 

0  =  Cells  of  the  nucleus  tubero-mammillaris  (nucleus  mammillo-lnfundib- 
ularis). 

•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 

[>  =  Cells  of  the  nuclei  tuberis  laterales   (n.  t.  1.).    (Examine  with  hand 

lens). 
O  =  Cells  of  the  ganglion  opticum  basale  (g.  o.  b.). 
Q  =  Cells  of  the  nucleus  ansae  peduncularis. 

°    =  Cells  of  the  ganglion  mediale  corporis  mammillaris. 

^  =  Cells  of  the  nucleus  intercalatus  corporis  mammillaris. 

.    =  Cells  of  the  cortex  cerebri. 

>-  =  Cells  of  the  globus  pallidus  nuclei  lentiformis. 


THE  JOHNS   HOPKINS   HOSPITAL   REPORTS. 


PLATE   IV. 


Fig.  11. 


Fig.  11a. 


Fig.  14. 


Fig.  14a. 


50  Edward  F.  Malonc. 


PLATE  V. 
Homo.    Series  D  (Co.ntixued). 

Magnification  of  Figs.  15  to  18  =  5  diameters. 
Magnification  of  Figs.  15A  to  ISA  =  2  diameters. 

f Cells  of  the  nucleus  tubero-mammillaris  (nucleus  mammillo  infun- 
0  =  .J      dibularis). 

I  Cells  of  the  nucleus  paraventricularis  hypothalami. 
•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 
A  =  Cells  of  the  nuclei  tuberis  laterales  (n.  t.  1.).     (Examine  with  hand 

lens). 
O  =  Cells  of  the  ganglion  opticum  basale  ( g.  o.  b. ). 
Q  =  Cells  of  the  nucleus  ansae  peduncularis. 
.    =  Cells  of  the  cortex  cerebri. 
>-  =  Cells  of  the  globus  pallidus  nuclei  lentiformis. 

Note  in  Fig.  17  the  closely  packed  group  of  cells  of  the  substantia  grisea 
situated  just  lateral  to  the  ventral  tip  of  the  third  vtntricle  (see  text). 

Note  in  Fig.  IS  the  cells  of  the  basal  optic  ganglion  crossing  mid  line 
(see  text). 


THE   JOHNS   HOPKINS  HOSPITAL   REPORTS. 


PLATE  V. 


Fig.  15. 


Fig.  ].5a. 


Fig.  16. 


Fig.  16a. 


Fig.  17. 


Fig.  17a. 


Fig.  18. 


Fig.  18a. 


52  Edward  F.  Malone. 


PLATE  VI. 

Macacus  Rhesus.    Sebies  A. 

Magnification  of  Figs.  19  to  22  =  6%  diameters. 
Magnification  of  Figs.  19A  to  22A  =  2V4  diameters. 

#  =  Cells  of  the  nucleus  tubero-mammillaris  (nucleus  mammillo-infundib- 
ularis). 

•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 
A  =  Cells  of  the  nuclei  tuberis  laterales   (n.  t.  I.). 
O  =  Cells  of  the  ganglion  opticum  basale   (g.  o.  b. ). 
JCells  of  the  corpus  hypothalamicum  (Luysil). 
Q~  \  Cells  of  the  nucleus  ansae  peduncularis. 

°    =  Cells  of  the  ganglion  mediale  corporis  mammillaris. 

^  =  Cells  of  the  nucleus  intercalatus  corporis  mammillaris. 

.    =  Cells  of  the  cortex  cerebri. 

>-  =  Cells  of  the  globus  pallidus  nuclei  lentiformis. 

Note  in  Fig.  22  the  group  of  densely  packed  cells  of  the  substantia  grisea 
located  just  lateral  to  the  third  ventricle   (see  text). 


THE  JOHNS  HOPKINS  HOSPITAL   REPORTS. 


PLATE  VI. 


Fig.  19. 


FiQ.  19a. 


'■v;m 


Fig.  20. 


Fig.  20a. 


Fig.  22. 


Fig.  22a. 


54  Edward  F.  M alone. 


PLATE  VII. 

Macacus  Rhesus.     Sebies  A   (Continxied). 

Magnification  of  Figs.  23  to  25  =  6%  diameters. 
Magnification  of  Figs.  23A  to  25A  =  2%  diameters. 

{Cells  of  the  nucleus  tubero-mammillaris  (nucleus  mammillo-infun- 
dibularis). 
Cells  of  the  nucleus  paraventricularis  hypothalami. 
•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 
O  =  Cells  of  the  ganglion  opticum  basale  (g.  o.  b. ). 
0  =  Cells  of  the  nucleus  ansae  peduncularis  (ganglion  basale  of  Kolliker). 
.    =  Cells  of  the  cortex  cerebri. 
>-  =  Cells  of  the  globus  pallidus  nuclei  lentiformis. 

Note  in  Fig.  24  lateral  from  the  third  ventricle  the  densely  packed  group 
of  cells  of  the  substantia  grisea;  such  masses  of  the  substantia  grisea  should 
not  be  confused  with  the  nuclei  tuberis  laterales. 


THE  JOHNS   HOPKINS  HOSPITAL   REPORTS. 


PLATE  VII. 


Fig.  2a 


Fig.  23a. 


.W 


FiC.  24. 


9,  S      •    .-' 


'^'*«??o-  *    *-  .':•'.--•-":'■*•••• 

'i^ct"";.-^  ;.:.:.•..■.■:■••.■.•.  v. 


Fig.  25. 


FlG.    20A. 


56  Edward  F.  Malone. 


PLATE  VIII. 
Lemur  Rdfus.     Series  A. 

Magnification  of  Figs.  26  to  30  =  SVa  diameters. 
Magnification  of  Figs.  26A  to  30A  =  3%  diameters. 

rCells  of  the  nucleus  tubero-mammillaris  (nucleus  mammillo-infun- 
0  =  -i      dibularis). 

I  Cells  of  the  nucleus  paraventricularis  hypothalami. 
•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 
O  =  Cells  of  the  ganglion  opticum  basale   (g.  o.  b.). 
>-  =  Cells  of  the  globus  pallidus  nuclei  lentiformis. 
Note  in  Fig.  27  the  absence  of  the  basal  optic  ganglion  (see  text). 


THE  JOHNS   HOPKINS  HOSPITAL   REPORTS. 


PLATE   VIII. 


Fig.  27. 


.    •     ■    •     • 
.     .  ■  ••••••■ 

if*-'--'       .*•»       " 

•.•;.•••■»'.■••.•• 


v.\    /■.•■.•.•:•■.'••;■  ■  ■     .  •    * 

•••I    I.  .••.•:.■■.••••  ••  ■  *  > 

"•.■:\  /.••■•.v-.v.^.--..--.  •  »£,•;  .°^o  •    • 


Fig.  28. 


t!t  .■••.•••  ♦. 


■:::•.■.•:■:,■■»■  ■ 


■'■■  r' 


Fig.  29. 


Fig.  30. 


Fig.  2Sa. 


Nuc\e<(5   Piraverttr'ic 
u\*r>5  liyPoth»Uw'\ 


optiCo-m 


©■fi 


T^- 


C  h  iASITiA 


58  Edward  F.  Malone. 


PLATE  IX. 

Cat.    Series  C. 

Magnification  of  Figs.  31  to  35  =  Sy^  diameters. 
Magnification  of  Figs.  31A  to  35A  =:  3';;  diameters. 

(Cells  of  the  nucleus  tubero-mammillaris  (nucleus  mammillolufun- 
dibularis). 
Cells  of  the  nucleus  paraventricularis  hypothalami. 
•  =  Cells  of  the  substantia  grisea  ventriculi  tertii. 
O  =  Cells  of  the  ganglion  opticum  basale   (g.  o.  b.). 
.    =  Cells  of  the  cortex  cerebri. 
)-  =  Cells  of  the  globus  pallidus  nuclei  lentiformis. 


THE 


JOHNS   HOPKINS  HOSPITAL   REPORTS 
■••.»•     •  •  •    . 


:-;•■>•  •••■   •    .T    •  •  •  •  Vv-  •  .■ 


PLATE  IX. 


'  *  • 


■••;•.•••>••"•*.♦• 
••■•'.>■••    i*  •  •  • 


<  .• 


P«M--:  -..•'  .4K-:S> 


Fig.  32. 


l»»:.v«V4:«.— 


/•.••■•:••••••.• 

»•••: •■•:••'■••  v".     ■>. »    .         »  ■* 
l^•■•^^•:      •".".•■.•  •  *•    *  *■'.  ' 


»      Fig.  34a 


60  Edward  F.  M alone. 


PLATE  X. 
Homo.   SsatLES  AC.    Nuclei  Tubesis  Laterales. 

Fig.  36  represents  the  actual  appearance  of  a  portion  of  the  preparation 
from  which  Fig.  4  (PI.  I)  was  drawn.  The  preparation  was  projected  at 
a  magnification  of  83%  diameters  and  every  cell  outlined  in  pencil.  Then 
under  the  microscope  each  cell  (as  far  as  this  was  possible)  was  identified 
and  reproduced  in  water  color  exactly  as  it  appeared.  This  Illustration 
is  accordingly  an  accurate  reproduction  of  the  appearance  of  the  preparation 
except  that  bloodvessels  and  neuroglia  have  been  omitted. 

Magnification    (after   y^    reduction)  =  55%   diameters. 

Comparing  Fig.  36  with  Figs.  4  and  4A  it  is  evident  that  the  region  illus- 
trated is  the  ventro-lateral  portion  of  the  tuber  cinereum.  At  the  right 
upper  corner  is  seen  a  portion  of  the  pes  pedunculi,  while  at  the  left  upper 
corner  appears  a  portion  of  the  fornix  column. 

The  two  groups  of  small  pigmented  cells  are  the  nuclei  tuberis  laterales. 
Note  that  they  are  partly  circumscribed  by  a  zone  containing  few  cells. 

The  large  cells  are  those  of  the  nucleus  tubero-mammillaris  (nucleus 
mammillo-intundibularis ) . 

The  small  pale  blue  cells  are  those  of  the  substantia  grisea  ventrlculi 
tertii.  Note  transition  types  between  these  cells  and  the  large  cells  of  the 
nucleus  tubero-mammillaris;  this  transition  involves  both  the  size  of  the 
cells  and  the  intensity  of  the  stain,  so  that  some  cells  occur  which  could  not 
be  positively  assigned  to  either  of  these  two  cell  groups. 

Compare  the  various  cell  groups  of  Fig.  36  with  the  corresponding  cell 
types  of  PI.  XII,  where  the  cells   (man)  are  highly  magnified. 


JOHNS   HOPKINS   HOS 


VOLUME    XVil. 


JOHNS  HOPKINS  HOSPITAL  REPORTS. 


VOLUME    XVII 


/^ 


( 


C 


62  Edward  F.  M alone. 


PLATE  XI. 
Homo.   Series  AC.    Ganglion  Optkum  Basale. 

Fig.  37  shows  the  actual  appearance  of  a  portion  of  the  basal  optic  gang- 
lion in  the  preparation  from  which  Fig.  9  (Plate  III)  was  drawn.  As  in  the 
case  of  Fig.  36  the  preparation  was  projected  (but  at  a  magnification  of 
145  diameters  instead  of  S3%)  and  every  cell  outlined  in  pencil.  Then 
under  the  microscope  every  cell  (except  in  rare  instances  where  this  was 
impossible)  was  identified  and  reproduced  accurately  in  water  color. 

Magnification  (after  %  reduction)  =  96%  diameters. 

Fig.  37  represents  the  whole  breadth  of  the  basal  optic  ganglion  near  its 
medial  pole  (Fig.  9,  PI.  Ill),  and  shows  the  relation  of  this  nucleus  to  the 
substantia  grisea  of  the  third  ventricle.  To  the  right  is  seen  the  ventro- 
lateral surface  of  the  brain  (anterior  perforated  substance). 

The  large  deeply  staining  cells  are  those  of  the  basal  optic  ganglion. 
Note  that  it  is  sharply  separated  from  the  substantia  grisea.  The  long 
coarse  processes  of  the  cells  of  the  basal  optic  ganglion  are  here  represented 
as  blue,  but  in  reality  they  are  almost  colorless;  this  figure,  however,  repre- 
sents correctly  the  peculiar  appearance  of  an  intercellular  feltwork,  to 
which  these  processes  give  rise. 

The  small  pale  cells  are  those  of  the  substantia  grisea  ventriculi  tertii. 
Note  the  difference  in  appearance  of  the  intercellular  substance  from  that 
within  the  basal  optic  ganglion,  due  to  the  absence  of  coarse  cell  processes. 

As  in  Fig.  36  bloodvessels  and  neuroglia  have  not  been  drawn. 

See  PI.  XII  for  the  appearance  of  these  two  cell  types  (in  man)  under 
higher  magnification. 


JOHNS  HOPKINS   HOS 


VOLUME   XVII. 


JOHNS  HOPKINS  HOSPITAL  REPORTS. 


VOLUME  XVII, 


'•    •    * 


^ 


'    1  i 


•  e 


\« 


^'^ 


1*   ' 


.  v•.,,»>,.v^\  v.-  -: 


v,» 


i   I 


C) 


64  Edward  F.  Malone. 


PLATE  XII. 
Homo.     MAOiNiFicAxio.N  =  580   Diameters. 

38.  Cells  of  the  ganglion  opticum  basale. 

39.  Cells  of  the  nuclei  tuberis  laterales. 

40.  Cells  of  the  substantia  grisea  ventriculi  tertii. 

41.  Cells  of  the  nucleus  tubero-niammillaris   (formerly  known  as  nucleus 

mammillo-infundibularis). 

42.  Cells  of  the  nucleus  ansae  peduncularis  (also  known  as  ganglion  basale 

of  Kolliker). 

43.  Cells  of  the  nucleus   paraventricularis   hypothalami. 


JOHNS  HOPKINS  HOSPITAL  REPORTS. 


VOLUME   XVII. 


38 


41 


■^» 


42 


39 


fes^ 


40 


•) 


66  Edward  F.  Malone. 


PLATE  XIII. 

Macacus  Rhesus.    Magnification  =  580  Diameters. 

44.  Cells  of  the  ganglion  opticum  basale. 

45.  Cells  of  the  nuclei  tuberis  laterales. 

46.  Cells  of  the  substantia  grisea  ventriculi  tertii. 

47.  Cells  of  the  nucleus  tubero-mammillaris   (formerly  known  as  nucleus 

mammillo-infundibularis). 

48.  Cells  of  the  nucleus  ansae  peduncularis   (also  known  as  the  ganglion 

basale  of  Kolliker). 

49.  Cells  of  the  nucleus   paraventricularis   hypothalami. 


JOHNS  HOPKINS  HOSPITAL  REPORTS. 


VOLUME   XVII. 


44 


47 


:    .    .      •  it-- 


J'-^--^ 

V,'-  \ 


48 


45 


^ 


t^'- 


46 


49 


wm 


,^ 


r. 


i 


68  '  Edward  F.  Malone. 


PLATE  XIV. 

Lemur  Rufus.    Maomficatiox  =  5S0  Diametebs. 

50.  Cells  of  the  ganglion  opticum  basale. 

51.  Cells  of  the  substantia  grisea  ventriculi  tertii. 

52.  Cells  of  the  nucleus    tubero-mammillaris     (formerly    known    as    the 

nucleus  mammillo-infundibularis ) . 

53.  Cells  of  the  nucleus  ansae  peduncularis   (also  known  as  the  ganglion 

basale  of  Kolliker). 

54.  Cells  of  the  nucleus   paraventricularis   hypothalami. 


JOHNS   HOPKINS   HOSPITAL  REPORTS. 


VOLUME   XVI 


50 


"^^ 


52 


53 


*   J^J^ 


r*\ 


"■^ 


f  <   * 


.^1 


54 


51 


-i) 


'^ 


70  Edward  F.  Malone. 


PLATE  XV. 
Cat.    Magnification  —  5S0  Dia.metebs. 

55.  Cells  of  the  ganglion  opticum  basale. 

56.  Cells  of  the  substantia  grisea  ventriculi  tertii. 

57.  Cells  of  the  nucleus    tubero-manimillaris     (formerly    known    as    the 

nucleus  mammillo-intundibularis). 
5S.  Cells  of  the  nucleus   paraventricularis   hypothalami. 


JOHNS  HOPKINS   HOSPITAL  REPORTS. 


VOLUME   XVII. 


55 


ilC 


iW^" 


56 


57 


58 


^   ^ 


SEPARATE  MONOGRAPHS  REPRINTED  FROM  THE  JOHNS 
HOPKINS  HOSPITAL  REPORTS. 

Studies  in  Bermatology.  By  T.  C.  Gilchrist,  M.  D.,  and  Emmet  Rixfobd, 
M.  D.    164  pages  and  41  plates.    Price,  In  paper,  $3.00. 

The  Malarial  Fevers  of  Baltimore.  By  W.  S.  Thayer,  M.  D.,  and  J.  HEn/yET- 
SON,  M.  D.  And  A  Study  of  some  Fatal  Cases  of  Ualaria.  By  Leweixts 
F.  Barker.  M.  B.     280  pages.     Price,  in  paper,  $2.75. 

The  Pathology  of  Toxalbnmln  Intoxications.  By  Simon  Plexnee,  M.  D. 
150  pages  with  4  lithographs.     Price,  In  paper,  $2.00. 

Studies  in  Typhoid  Fever,  I,  II,  III.  By  William  Oslee,  M.  D.,  and  others. 
Extracted  from  Vols.  IV,  V  and  VIII  of  the  Reports.  757  pages. 
Price,  in  cloth,  $5.00. 


Pneumothorax.    A    Historical,    Clinical,    and    Experimental    Study. 
Charles  P.  Emebson,  M.  D.    Price,  in  paper,  $4.00. 


By 


NEW  SERIES. 

I.  Free  Thrombi  and  Ball-Thrombi  in  the  Heart.    By  J.  H.  Hewitt,  M.  D. 
82  pages  and  20  illustrations.    Price,  in  paper,  $1.00. 

II.  Benzol  as  a  Leucotoxin.  By  Laurence  Selling,  M.  D.  60  pages  and 
3  plates,  1  in  colors.    Price,  in  paper,  $1.00. 

III.  Primary  Carcinoma  of  the  Liver.  By  M.  C.  Winteenitz,  M.  D.  42 
pages.     Price,   in   paper,   75   cents. 

IV.  The  Statistical  Experience  Data  of  The  Johns  Hopkins  Hospital,  Balti- 
more, Md.,  1892-1911.  By  Fkkuerick  L.  Hoffmak,  LL.  D.,  F.  S.  S.  IGl 
pages.    Price,  in  paper,  $2.00. 

V.  The  Origin  and  Development  of  the  Lymphatic  System.  By  Florence  R. 
Sabik.    94  pages.    Price,  in  paper,  $2.00. 

VI.  The  Nuclei  Tuberis  Laterales  and  the  So-called  Ganglion  Opticum  Basale. 
By  EmvAiii)  F.  Malone.    70  pages  and  l.j  plates.    Price,  in  paper,  $1.00. 


UNIVERSITY  OF  CALIFORNIA  LIBRARY 

Los  Angeles 
This  book  is  DUE  on  the  last  date  stamped  below. 

OCT  2  2  ]QR2 
OCT  16  RfflT 

,UG  1  2  '''' 

BlOMti^  UB. 

TiOV  17K£<« 

flAR2  4188l 

B^  NOV  19  ^969 

8ir),v--^_   ,j^_ 

biomedMAY^TB 

r 

- 

Torm  Ij9-50»b-11,'o0  (2554)4 

44 

T^iir  •  - 

UNIVERSITY  OF  LA1.IFORNU 


D     000  117  154     5 


ii 


